Florissantoraphidia, Makarkin, Vladimir N. & Archibald, S. Bruce, 2014

Makarkin, Vladimir N. & Archibald, S. Bruce, 2014, A revision of the late Eocene snakeflies (Raphidioptera) of the Florissant Formation, Colorado, with special reference to the wing venation of the Raphidiomorpha, Zootaxa 3784 (4), pp. 401-444 : 412-414

Family Raphidiidae Latreille, 1810

Diagnostic character states of venation. Forewing: In subcostal space, two crossveins: one connecting ScP, RA; other closing pterostigma proximally. At least one branch of RA incorporating in pterostigma. In RA spaces, two crossveins between RA, RP (three crossveins in Mongoloraphidia abnormis Liu et al., 2010a : Fig. 1 ). M, CuA appear to originate from R either separately or at same point (rarely stem M+CuA present). 1r-m connects RP and MA. In medial space, only two crossveins between MA, MP forming two doi. CuA fused with MP for short distance. AA2, AA3 basally almost entirely fused.

Hind wing: In subcostal space, one crossvein closing pterostigma proximally; at least one branch of RA incorporating in pterostigma; two doi (except normally one in few species); CuP, AA1 medially fused (except two Baltic amber species).

Composition. Twenty six extant genera (ca. 200 species) distributed in the Northern Hemisphere (Aspöck et al. 2012a) and four fossil genera from the late Eocene of Baltic amber (Succinoraphidia Aspöck et Aspöck, 2004) and Florissant ( Megaraphidia Cockerell, 1907 ; Archiraphidia ; Florissantoraphidia gen. nov.).

Besides seven from Florissant, the following six named species have been described: ‘ Agulla ’ protomaculata Engel, 2011 (late Ypresian of the Green River Formation, Parachute Creek Member of Colorado); ‘ Raphidia ’ baltica and Succinoraphidia exhibens (late Eocene of Baltic amber); ‘ Raphidia ’ creedei (late Oligocene of Creede, Colorado); Agulla mineralensis (middle Miocene of Stewart Valley, Nevada); and ‘ Ohmella ’ coffini (late Miocene of Ardèche (Mt. Andance), France). Also, six specimens have been figured, but not named, from the early Oligocene of the Anna potash mine in Haut-Rhin department, France ( Nel 1993: Fig. 10 ); the late Oligocene of Enspel, Germany ( Wedmann 2000: Fig. 13 ); the early Miocene of Rubielos de Mora, Spain ( Peñalver & Seilacher 1995: Fig. 8 ), and the late Miocene of Murat, France ( Nel 1993: Figs 6–9 ).

Comments on characters. The additional distal crossvein between C/ScP and RA (2 b scp-r). This crossvein is absent in extant Raphidiidae , except in the three species (both in fore- and hind wings) of the subgenus Harraphidia Steinmann, 1963 of the genus Harraphidia , distributed in Morocco and extreme south of Spain ( Monserrat & Papenberg 2006: Fig. 1 ). 2b scp-r is present in at least one Florissant species, Archiraphidia tumulata ( Scudder, 1890) . Carpenter (1936) identified this crossvein in one forewing of the ‘ Raphidia’ creedei holotype, but found it certainly absent in the other.

Discoidal cells (doi). Some specimens of species which normally have two doi may adventitiously possess other numbers of these. Figured examples of such aberrant individuals include an instance of one doi in Mongoloraphidia kaszabi Aspöck et Aspöck, 1968 ( Aspöck & Aspöck 1968a: Fig. 1 ), Alena infundibulata Aspöck et al., 1994 : Fig. 18 , and Raphidia ophiopsis Linnaeus, 1758 ( Aspöck et al. 1977: Fig. 1 e); three in Turcoraphidia acerba ( Aspöck & Aspöck, 1966) ( Aspock et al. 1991: Fig. 18 ); and four in Raphidia ophiopsis ( Aspöck et al. 1977: Fig. 1 h). One doi is characteristic of all species of the genus Harraphidia ( Monserrat & Papenberg 2006) (see below).

Remarks. No raphidiopteran specimens with the full combination of the raphidiid diagnostic character states in their venation are known before the Eocene, except probably one mesoraphidiid forewing (see above; but this is incompletely preserved). The two genera described from the Middle Jurassic of Daohugou ( Engel & Ren 2008) were placed in Raphidiidae ( Ren et al. 2010) . However, they cannot be assigned to this family, as both genera lack a crossvein closing the pterostigma proximally, and the third doi is present.

No venational autapomorphies of Raphidiidae were identified by Aspöck and Aspöck (2004); however, we find that MP fused for a short distance with CuA in the forewing may be considered one. The CuA fused for some distance with AA 1 in the hind wing is an apomorphy characterisizing all extant raphidiids. However, this condition probably appeared at later stage; it is absent in the two Baltic amber species (above).

Generic attribution of Florissant raphidiid species. Extant genera and species of Raphidioptera are based on genitalic characters, as eidonomic characters (external, visible without dissection) have been widely considered too conservative to be useful at those levels (e.g., Aspöck 1986). This has rendered the fossil history of the order obscured below the family level.

Florissant species are currently assigned to the extant genus Raphidia , a situation that most recent authors recognize as a problematic lumping (e.g., Aspöck et al. 1991; Engel 2002). Alternatively, Engel (2002) suggested that all North American fossil Raphidiidae might be assigned to Agulla , citing lack of reliable wing characters with which to confidently separate them further.

We examined wing venation within the order, and found broad patterns that covary with genera or generic groups that are congruent with those that have been independently based on genetalic characters, and so appear useful in distinguishing the generic relationships of extinct species known only or primarily by their wings. Based on these characters, described in the genus diagnoses provided here, we found three distinct species groups of Florissant raphidiid species which we recognize as the genera Megaraphidia , Archiraphidia , and Florissantoraphidia gen. nov.

All three of these genera share a combination of the proximal location of 3ra-rp (located within the pterostigma) in both their fore- and hind wings and the presence of the long basal 1r-m in the hind wing. Both conditions are plesiomorphic in the order, which precludes their usefulness in assessing its internal relationships. In extant snakeflies, this combination is present in some species of the raphidiid genera Alena , Agulla , Dichrostigma , and Phaeostigma . In these cases, however, their venation differs from the Florissant species in other aspects, suggesting that the Florissant species cannot be confidently attributed to these or other extant genera; they might be closely related. This placement of 3ra-rp excludes these fossil species from Raphidia , where, in all species this crossvein is almost invariably located distad the pterostigma, rendering the distal portion of RA zigzagged. We know of only a single specimen of Raphidia ophiopsis where 3ra-rp is aberrantly located within the pterostigma (Steinmann 1964: Fig. 26; as Raphidia helvetica Steinmann, 1964 ); all others that we are aware of have 3ra-rp located distad the pterostigma (see e.g., Aspöck et al. 1977: Figs 1 a–h). Venational character states indicate the following possible close relationships between Florissant and extant genera: Megaraphidia and Phaeostigma , Archiraphidia and Harraphidia , and Florissantoraphidia gen. nov. and Agulla or Alena .

We recognize that at least two of these three Florissant genera are preliminary; however, this status has more confidence than leaving them in Raphidia where they clearly do not belong, and which would render that genus paraphyletic.

Figures

FIGURE 1. Wing venation of the Jurassic Mesoraphidiidae. A, Mesoraphidia inaequalis Martynov, 1925 (Late Jurassic of Karatau, Kazakhstan), forewing. B, same, hind wing. Re-drawn with slight simplification from Martynov 1925 b. Fused veins are shown by red lines. Identity of the vein stub labeled “? ” in basal-most part of forewing is unknown. Both wings are shown with apex to right. No scale provided or pterostigma shown in original figure.

FIGURE 6. Megaraphidia exhumata (Cockerell, 1909). Holotype UCM 4513. A, the specimen as preserved. B, forewing venation. Scale bar = 4 mm (both to scale).

FIGURE 7. Megaraphidia exhumata (Cockerell, 1909). Holotype MCZ 3640. A, part (converted to right). B, counterpart. C, forewing venation. Scale bar = 4 mm (all to scale).

FIGURE 8. Megaraphidia sp. Specimen MCZ 3641. A, part (converted to right). B, counterpart. C, hind wing venation. Scale bar = 4 mm (all to scale).

FIGURE 9. Pterostigma of Archiraphidia species. A – C, A. tumulata (Scudder, 1890): A, left wings. B, right forewing. C, right hind wing. D – F, A. tranquilla (Scudder, 1890): D, four wings of the holotype MCZ 446. E, four wings of MCZ 4137 (lectotype of A. eventa (Scudder, 1890 )); F, left wings of MCZ 253 (paralectotype of A. eventa). G, three wings of A.? somnolenta (Scudder, 1890). H, right forewing of A.? somnolenta. HW, hind wing; LFW, left forewing; RFW, right forewing. All wings are shown with apex to right and not to scale.

FIGURE 10. Archiraphidia tumulata (Scudder, 1890). Holotype MCZ 244. A, counterpart. B, part (converted to right). Scale bar = 4 mm (both to scale).

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).

FIGURE 18. Archiraphidia? somnolenta (Scudder, 1890). Wing venation of the holotype MCZ 488. A, left forewing. B, right forewing as preserved. C, right forewing, reconstruction. D, left hind wing. E, right hind wing. Scale bar = 4 mm (all to scale).