Nobloedischia rasnitsyni, a new genus and species of Oedischiidae (Orthoptera) from the Lower Permian Wellington Formation of Oklahoma, USA

Abstract Nobloedischia rasnitsyni gen. et sp. n. (Oedischiidae) is described from the Lower Permian Wellington Formation of Noble County, Oklahoma. The genus is similar to both Petrelcana (Oedischiidae: Mezenoedischiinae) and Oedischia (Oedischiidae: Oedischiinae) and is left unplaced at the subfamily level. The new species is the twelfth Orthoptera species and the fourth species of Oedischiidae from these deposits.

Of the more than 200 species of insects known from the Lower Permian Wellington Formation of Kansas and Oklahoma, USA (Beckemeyer and Hall 2007), eleven are in the order Orthoptera: eight from the Elmo, Kansas deposits (Tillyard 1932;1937;Carpenter 1943Carpenter , 1966Gorochov 1987), and three from the Midco, Oklahoma (Noble County) formations. Four of the eleven species, one from Kansas and three from Oklahoma, were described only recently (Béthoux and Beckemeyer 2007;Beckemeyer 2011). The new taxon described here constitutes the tenth genus and twelfth species of Wellington Formation Orthoptera (see Beckemeyer 2011 for a list of previously described species). It seems most closely related to Oedischia and Petrelcana.

Materials and methods
This description is based on a forewing specimen (Figs 1, 2) collected by Don Arnold and Rick Grantham of Oklahoma State University (OSU). The holotype is in the K. C. Emerson Museum, Department of Plant Pathology and Entomology, OSU. The specimen comprises part and counterpart of a nearly complete wing, absent approximately the basal 20% of the wing length (including the precostal area), the anal field, and the posterior margin of the cubital field; cross veins are poorly preserved in the distal radial and medial fields.
Photomicrographs of the part and counterpart were made using a Nikon 990 digital camera with an American Optical dissecting microscope and an external strobe flash (Nikon SB-26). Flash orientation was optimized to show details of interest. The photographed images were processed using Adobe Photoshop 6.0 and imported into a vector-graphics software program (XARA Extreme 4.0, XARA Group, Ltd., London), where they were assembled into composite images. The venation reconstruction drawing was made as an overlay (Fig. 3).
Venation interpretation and notation follow the system of Nel (2001, 2002). Since that system is relatively new, not uniformly accepted, and not in as wide use as the terminology of Sharov (1971), Gorochov (1987Gorochov ( , 1995, Carpenter (1992), and Gorochov and Rasnitsyn (2002), I list here the relationship between the most common notation systems: Diagnosis. Nobloedischia is differentiated from Oedischia by branches of RA directed distally, first branching of RA at level of fusion of MA1 and RP; fusion of RP and MA1 (the anterior branch of MA) long rather than limited to a point of contact; field between MP and CuA+CuPaα narrower than area between MP and MA/MA2 at least in basal 2/3 of length; CuA+CuPaα and its branches reach posterior wing margin obliquely.

Béthoux and Nel Sharov/ Gorochov Carpenter Gorochov and Rasnitsyn
Differs from Petrelcana by width of costal and subcostal fields at level of branching of M about equal rather than costal field much wider than subcostal field; cross veins between RA and RP without secondary cross veins; branching of MA moderately remote from anastamosis with RP (as in Gorochov 1995, Fig. 142) rather than MA branching close to fusion with RP, stem of MA1 before fusion an obvious branch of the longitudinal vein rather than this segment of MA1 very short and appearing as a thickened cross vein between MA/MA2 and RP+MA1 (as in Gorochov 1995, Fig. 144; stems of CuA and CuPaα about equal rather than stem of CuA very short.  1) and 826b, counterpart (Fig. 2), comprising the only Orthoptera fossil on slabs densely covered with multiple insect wing fragments of varied orders and with Conchostraca fossils as well. The slab containing 826a is approximately 12 by 21 cm and contains a total of 88 insect specimens (818a through 905a); the second slab (8 by 17 cm) contains 78 insect specimens (818b through 896b).

Nobloedischia rasnitsyni
Etymology. The specific epithet, rasnitsyni, is an honorific for eminent scientist Dr. Alexandr P. Rasnitsyn in recognition of his long and productive career, and his invaluable and varied contributions to paleoentomology.
Diagnosis. As for genus. Description. Forewing. Preserved length 32 mm, estimated length 39 mm (basal 20%, including precostal area, missing); width 7 mm. ScP terminates at 78% wing length, cross veins simple, moderately spaced, and oriented normal to costal margin and ScP, maximum width of costal field 1.4 mm, maximum width of subcostal field 1.0 mm, ratio width of subcostal to width of costal field 1.4:1. RA sigmoidal in form, bending anteriorly at level of separation of Cu and M (35% of wing length), bending posteriorly at level of fusion of MA and RP, where RA branches; RA branching anteriorly pectinate, six branches reaching wing margin, basal branches intersecting margin obliquely, distal branches nearly longitudinal in orientation; posterior-most branch of RA terminating at 95% wing length; cross veins between ScP and RA similar in form and spacing to those between ScP and costal margin. RP origin at 51% wing length; RP fusion with MA1 at 62% of wing length, length of fusion 0.9 mm; RP with four posteriorly pectinate branches, simple or distally twigged, first branch at 72% wing length, anterior-most branch straight and terminating near apex of wing at 98% wing length; cross veins between RA and RP with veins simple, some bowed with center of curvature basal. M+CuA branches at 35%, M branches at 37%, length of M from separation from M+CuA to branch 0.9 mm, MA branches at 59%, MA2 branches at 69% of wing length each branch with terminal twig, MA1 apparently simple, terminating at 89%, free length of MA1 before fusion with RP 1.0 mm. MP distally twigged, terminating on posterior margin at 74% and 77% wing length. CuA+CuPaα with four branches; the basal branches of CuA+CuPaα intersect the posterior margin obliquely, with the angle between the branches and the wing margin increasing distally (angles of intersection are 12°, 13°, 23°, and 32°, for the successive branches); CuPa branches at 34% of the wing length, free length of CuPaα before fusion with CuA 1.1 mm, length of CuA from separation from M+CuA to fusion with CuPaα 0.9 mm, CuPaβ straight as preserved, cross veins between CuPaβ and CuA+CuPaα normal to the longitudinal veins, becoming oblique and oriented longitudinally after the first branching of CuA+CuPaα; short segment of CuPb that is preserved (extending from 20% to 35% of wing length) is straight; anal veins not preserved. Figures 1-3. Nobloedischia rasnitsyni gen. et sp. n. 1. Holotype specimen, part, OSU826a. Forewing in dorsal aspect 2. Holotype specimen, counterpart, OSU826b. Forewing in ventral aspect 3. Venation reconstruction and notation following  Photographs and drawing to same scale; scale bar 5 mm.

Discussion
The combination of character states of N. rasnitsyni make placement in one of Gorochov's subfamilies somewhat problematic; it would perhaps come closest to fitting in Oedischiinae. Nobloedischia shares with Oedischia the characters: origin of anterior branch of MA moderately remote from its anastamosis with RP (as in Gorochov 1995: Fig. 142); distance from branching of M+CuA to branching of M subequal to distance from branching of M+CuA to fusion of CuA and CuPaα, and greater than length of cross veins between CuA+CuPaα and MP; ratio of width of costal field (anterior margin-ScP) to width of subcostal field (ScP-RA) less than 1.5; field between RA and RP with some cross veins bowed and/or sigmoidal rather than straight; CuPaα medium sized basal of its fusion with CuA (last three characters were used by , to distinguish the clade (Gerarus bruesi Meunier, 1909 + O. williamsoni)). On the other hand, Nobloedischia is differentiated from Oedischia by the characters listed in the Diagnosis (vide supra).  used a total of 74 characters in their cladistic phylogenetic analysis of the Orthoptera; I have not repeated that analysis, but did code the characters and found that of the 61 character states that could be compared (the remaining characters were not preserved in available specimens of one or the other of the taxa) between Nobloedischia and Oedischia, 52 matched and 9 did not.
For the other oedischiids, the next closest pairing was between Nobloedischia and Elcanoedischia, with 45 of 58 character states matching. However, Nobloedischia differs from genera in Elcanoedischiinae Gorochov 1987 by branching of MA moderately remote from anastamosis with RP (as in Gorochov 1995: Fig. 142) rather than MA branching close to fusion with RP, with this short segment of MA appearing as a thick cross vein between MA/MA2 and RP+MA1 (as in Gorochov 1995: Fig. 144). Nobloedischia can be differentiated from genera in Tettoedischiinae Gorochov, 1987 by costal field not greatly wider than subcostal field; base of MP well basal rather than at or distal to base of RP; branching of CuPb basal rather than distal to branching of M+CuA. Nobloedischia differs from genera in Mezenoedischiinae by the features listed in the Diagnosis section for Petrelcana (vide supra). Because of the presence of these conflicting character states, at this point I prefer to leave the subfamilial assignment undetermined.
The high number of densely distributed specimens (wings and wing fragments) present on the slabs containing this species seems to support Hall's (2004) thesis that the Oklahoma Wellington Formation insect deposit facies are derived from marginal marine lagoon sediments, with insect remains likely comprised of allochthonous material washed in by streams.
chael Engel sponsors my work through my affiliation with his Paleoentomology Laboratory. Three anonymous reviewers and the editor, Dr. Dmitry Shcherbakov, made valuable and constructive comments that significantly improved this paper. The work reported here is a contribution of the University of Kansas Natural History Museum Division of Entomology.