Parabintoniella papierae n. gen., n. sp., a new Triassic representative of the Bintoniellidae, a poorly known extinct family of Orthoptera (Insecta) Author Béthoux, Olivier Author Grauvogel-Stamm, Léa text Geodiversitas 2022 2022-06-23 44 21 603 608 journal article 86005 10.5252/geodiversitas2022v44a21 b760c638-30c3-40d1-9c3f-d3cc43476b53 1638-9395 6787056 urn:lsid:zoobank.org:pub:1928AA5E-1D54-434A-829F-68DB923EBF1B Parabintoniella papierae n. gen., n. sp. ( Figs 1 ; 2 ) urn:lsid:zoobank.org:act: 1AAFA563-F408-41C0-A519-B1E25EC3ABCA HOLOTYPE . — Specimen 5589/5590 (collection Louis Grauvogel at the ‘Staatlisches Museum für Naturkunde Stuttgart’, Stuttgart, Germany) ( Fig. 1A, B ). PARATYPE . — Specimen 9122a, b (collection Louis Grauvogel at the ‘Staatlisches Museum für Naturkunde Stuttgart’, Stuttgart, Germany) ( Figs 1C, D ; 2 ). DIAGNOSIS. — Forewing (and see Fig. 3 ): no distinct base of CuPaα diverging from CuPa (and fusing with CuA or M+CuA) (derived state shared with all Bintoniellidae as delimited herein); MA simple (derived state shared with Bintoniella brodiei and Eubintoniella ferganica ); CuA+CuPaα with more than two branches (plesiomorphy shared with Proshiellinae and Oshiellinae); cross-veins forming loose, convex intercalaries between ScP branches (shared with Proshiellinae , polarity unclear). ETYMOLOGY. — The species epithet honours the late Francine Papier, who significantly contributed to the knowledge of the ‘Grès à Voltzia’ fossil insects. TYPE LOCALITY AND STRATIGRAPHY. — The holotype specimen 5589/5590 was collected at Gottenhouse ( Bas-Rhin , France ) and the paratype specimen 9122a, b was collected at Bust ( Bas-Rhin , France ). Both specimens come from the ‘Grès à Voltzia’ (Upper Buntsandstein; early Middle Triassic; Gall & Grauvogel-Stamm 2005). GENERAL DESCRIPTION Forewing: length about 22.4-28.0 mm, width 6.4-8.0 mm; ScA conspicuous; in its median part, area between anterior wing margin and ScP broad, with cross-veins forming loose, convex intercalaries between ScP branches (concave); ScP reaching the anterior wing margin from two-thirds to threequarters of wing length; RA/RP split slightly distal to wing mid-length; RA with anterior veinlets and cross-veins between them; RP without well-organized branching pattern, with five to eight terminal branches/veinlets covering the apex; M+CuA(+CuPaα) splitting into M and CuA+CuPaα at about the first third of wing length; free basal portion of M (between the M/CuA(+CuPaα) split and the MA/MP split) relatively long; both MA and MP simple (known in holotype only); CuA+CuPaα posteriorly pectinate, with four branches; no distinct base of CuPaα diverging from CuPa (and fusing with CuA or M+CuA; better visible in specimen 9122); CuPaβ and CuPb simple; CuPb reaching posterior wing margin at wing mid-length; AA with at least four, simple veins; cross-veins reticulated (i.e. forming 2-3 cells between two main veins) between ScP veinlets, in the apical half, elsewhere mostly scalariform; along the posterior wing margin, areas between CuPb and AA branches with numerous, sigmoidal cross-veins; membrane uniformly hyaline. SPECIMEN DESCRIPTIONS Specimen 5589/5590 ( Fig. 1A, B ): adpression of an isolated, nearly complete left forewing, in both dorsal (5590) and ventral (5589) aspects, presumably of a female insect (see Remarks section); preserved length 26.6 mm (estimated complete length about 28.0 mm), width 8.0 mm (at its widest); RP with either eight terminal branches or seven with an intercalary vein between its most posterior branch and MA. Specimen 9122a,b ( Figs 1C, D ; 2 ): adpression of an isolated left forewing, in both dorsal (9122a) and ventral (9122b) aspects, with most of the postero-distal area and area basal to the claval furrow (itself located along CuPb; i.e. the remigulum) missing, presumably of a male insect (see Remarks section); preserved length 20.6 mm (estimated complete length about 22.4 mm ), width 6.4 mm . REMARKS From the difference in width of the two forewings at their widest, which is the single and most accurate comparable dimension between them, the size of the smallest one ( Fig. 1C, D ) is 80% that of the largest ( Fig. 1A, B ). This difference in size falls within the range of intra-specific variability and more particularly within that of sexual size dimorphism (SSD), commonly female-biased in extant Orthoptera ( Hochkirch & Gröning 2008 ) . Moreover, such phenomenon is also known to occur in early stem-Orthoptera ( Du et al. 2017 ) and has already been assumed for the related species Bintoniella brodiei , with a similar ratio ( Whalley 1982 ). As the other differences are minimal, it is doubtless that the two specimens belong to the same species, the specimen 5589/5590 presumably being a female, and the specimen 9122a, b a male. FIG. 1. — Parabintoniella papierae n. gen., n. sp. : A , B , holotype, specimen 5589/5590, left forewing (collection Louis Grauvogel at the ‘Staatlisches Museum für Naturkunde Stuttgart’); A , photograph, ventral aspect (5589), light-mirrored; B , drawing of venation (reconstructed parts in grey); C , D , specimen 9122a, b, right forewing (collection Louis Grauvogel at the ‘Staatlisches Museum für Naturkunde Stuttgart’); C , photograph, ventral aspect (9122b), light-mirrored (the white frame indicates the section enlarged in Fig. 2); D , drawing of venation (reconstructed parts in grey). Scale bar: 4 mm. FIG. 2. — Parabintoniella papierae n. gen., n. sp. , specimen 9122b (collection Louis Grauvogel at the ‘ Staatlisches Museum für Naturkunde Stuttgart’), photograph of detail of medio-cubital area (location as shown with a white frame in Fig.1C); the sign * indicates the area where the free basal portion of CuPaα, present in the Orthoptera groundplan but absent in the Bintoniellidae , could be expected to occur. Scale bar: 2 mm. With the aid of the systematic conventions posited above, the relationships between the species to which the new specimens belong can be discussed ( Fig. 3 ). The occurrence of the character state ‘in forewing, no distinct base of CuPaα diverging from CuPa (and fusing with CuA or M+CuA)’ (to be understood as ‘beyond the wing base’) qualifies an assignment to the Bintoniellidae . In other words, affinities with the Proshiellinae can be excluded. This condition is best observed in the specimen 9122 ( Fig. 2 ). FIG. 3. — Summary of the assumed phylogenetic relationships of the taxa and species considered in this contribution (character states invariant under different hypotheses in dark grey): A , favoured hypothesis, assuming that Parabintoniella papierae n. gen., n. sp. is the sister-group of Eubintoniella ferganica Gorochov, 1987 and Bintoniella brodiei Handlirsch, 1939 , owing to the occurrence of a simple MA in the forewing of these species; B , alternative hypothesis, assuming that Parabintoniella papierae n. gen., n. sp. is the sister-group of Oshiellinae, Eubintoniella ferganica and Bintoniella brodiei , owing to the occurrence of cross-veins between ScP veinlets in the forewing of Parabintoniella papierae n. gen., n. sp. In contrast to all the Proshiellinae and the Oshiellinae, in which MA commonly has 3-4 branches in the forewing, the species to which the new specimens belong displays a simple MA. This feature, which is shared with Bintoniella brodiei (see Whalley 1982 ; Béthoux 2012 ) and Eubintoniella ferganica (see the original description), is definitely derived and represents the diagnostic trait of a yet unnamed taxon. Furthermore, in contrast to Bintoniella brodiei and Eubintoniella ferganica , in which CuA+CuPaα has two branches (rare exceptions are known in Bintoniella brodiei , see Béthoux 2012 ), the species to which the new specimens belong displays a CuA+CuPaα with four branches. This trait must be regarded as a plesiomorphy, as Proshiellinae and Oshiellinae commonly possess a 4- to 5-branched CuA+CuPaα. It follows that the species to which the new specimens belong is the sister-group of Bintoniella brodiei and Eubintoniella ferganica ( Fig. 3A ). Unlike Bintoniellidae , in which no cross-veins occur between ScP branches, the species to which the new specimens belong displays cross-veins forming loose, convex intercalaries between ScP branches. This state occurring in Proshiellinae , it can then be regarded as a plesiomorphy within Bintoniellidae . Accordingly, the species to which the new specimens belong would be the sister-group of all other Bintoniellidae (namely, Oshiellinae, Bintoniella brodiei and Eubintoniella ferganica ; Fig. 3B ). Considering (i) that this scenario implies that the number of branches of MA would have been subjected to homoplasic evolution, and, on the other hand, (ii) that the occurrence of cross-veins between ScP veinlets, and their lack thereof, can both occur within a single genus (such as in the Permian stem-Orthoptera genus Tcholmanvissia Zalessky, 1929 ; see Béthoux & Nel 2002b ), we consider this hypothesis less likely.