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Wing Venation Morphology and Variability of Gerarus Fischeri (Brongniart, 1885) Sensu Burnham (Panorthoptera; Upper Carboniferou

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Wing Venation Morphology and Variability of Gerarus Fischeri (Brongniart, 1885) Sensu Burnham (Panorthoptera; Upper Carboniferou Org. Divers. Evol. 3, 173–183 (2003) © Urban & Fischer Verlag http://www.urbanfischer.de/journals/ode Wing venation morphology and variability of Gerarus fischeri (Brongniart, 1885) sensu Burnham (Panorthoptera; Upper Carboniferous, Commentry, France), with inferences on flight performance Olivier Béthoux1,2, André Nel1,* 1 Muséum National d’Histoire Naturelle, Paris, France 2 Botanique et Bioinformatique de l’Architecture des Plantes, CIRAD – CNRS, Montpellier, France Received 23 August 2002 · Accepted 4 March 2003 Abstract The Geraridae is a family with no consensual assignment. In order to improve this situation, a revision of the wing venation of Gerarus fischeri (Brongniart, 1885) sensu Burnham (1983) is presented. Gerarus bruesi Meunier, 1909, Archaeacridites elegantissima Meunier, 1909 and Sthenaropoda minor Handlirsch, 1919 are considered as new junior synonyms of G. fischeri.A new interpretation of the hind wing venation is pro- posed, yielding additional support for the ‘panorthopterid’ assignment.The wing venation of this species shows outstandingly high variability. For a better resolution of relationships of Geraridae within Panorthoptera, additional studies of representatives of the latter taxon are required. Key words: Panorthoptera, Geraridae, Gerarus fischeri,variability, wing venation Introduction erous fossils which likely belong to Gryllones [= Poly- neoptera of other authors]”. This definition summarizes The Geraridae has been a key-stone problematic family well the phylogenetic problem with this group. for a long time. Brongniart (1893), followed by Burn- In order to improve our knowledge of this family, prior ham (1983), Carpenter (1992) and Ross & Jarzembows- to a cladistic analysis of the Panorthoptera (Béthoux, in ki (1993), included it in a paraphyletic or even poly- prep.), we here present a revision of a species known phyletic Protorthoptera. Sharov (1961; 1968, translated from numerous and well-preserved specimens, Gerarus in 1971) considered it as the unique representative of the fischeri (Brongniart, 1885), known from the Upper Car- Protorthoptera, sister-group of Orthoptera. Kukalová- boniferous of Commentry (France). This revision is based Peck (1991) and Kukalová-Peck & Brauckmann (1992) on a set of specimen drawings instead of composite draw- advocated it as a representative of the ‘hemipteroid’ lin- ings, not given in previous publications. The study of eage, on the basis of a clypeal ‘bump’ tentatively inter- complete specimens and of all the available material re- preted as a suction pump. Béthoux & Nel (2002b: 8) de- veals a previously underestimated range of variation in nied any value to this character. Labandeira (1994) in- the wing venation of Gerarus fischeri. cluded the Geraridae in the Hypoperlida. Gorokhov (2001) considered it as a distinct order Gerarida, sister- group to the Titanoptera. Béthoux & Nel (2002a, b) con- Material and methods sidered it as closely related to Orthoptera, in the larger clade Panorthoptera. Rasnitsyn (2002) included the Ger- The venation patterns and vein widths were drawn with a aridae in the Eoblattida,“a mixture of stem (= ancestral stereo microscope and camera lucida directly from the fossil or close to ancestral) and insufficiently known Carbonif- surface, dry and under alcohol. Both print and counter-print *Corresponding author: André Nel, Laboratoire d’Entomologie and CNRS UMR 8569, Muséum National d’Histoire Naturelle, 45 rue Buffon, F-75005 Paris, France; e-mail: [email protected] 1439-6092/03/03/03-173 $ 15.00/0 Org. Divers. Evol. (2003) 3, 173–183 174 Béthoux & Nel were used when available. The drawings were readjusted on 1909d Archaeacridites elegantissima – Meunier, 146, photographs using image-editing software. text-fig. 17, pl. V, fig. 2. Photographs were taken using a digital camera, Nikon 1909d Sthenaropoda fischeri – Meunier, 148, text-figs Coolpix 990. The fossils were illuminated with fiber optics. 19–20, pl. V, figs 4, 4a. When possible, counter-prints were light-mirrored in order to reconstruct the relief in dorsal aspect. 1914 Archaeacridites bruesi – Meunier, 363 (citation). We follow the nomenclature of Béthoux & Nel (2002a, b) 1917 Oedischia fischeri – Lameere, 176–177 (citation, proposed for the ground plan of ‘panorthopteroid’ insects, but invalidated in the following). which is the most consistent with the organization of main 1917 Sthenaropoda fischeri – Lameere, 177–178 veins in Gerarus fischeri. (review). 1917 Sthenaropoda agnusi – Lameere, 178 (assign- ment of previous material in a new species, Systematic section original description). 1917 Sthenaropoda lerichei – Lameere, 178 (assign- Family Geraridae Scudder, 1885 ment of previous material in a new species, Genus Gerarus Scudder, 1885 original description). 1917 Archaeacridites bruesi – Lameere, 177, 178–179 Note on synonymy (review). Burnham (1983: 12) discussed the problem of doubts con- 1917 Archaeacridites elegantissima – Lameere, 179 cerning the priority of Scudder’s publication over Brong- (review). niart’s, both from 1885. Burnham ignored the date of pub- 1919 Sthenaropoda fischeri – Handlirsch, 548, text- lication of Brongniart’s work, but its front page indicates fig. 46, pl. 14, fig. 14. that it was published within the first semester of the year 1919 Sthenaropoda bruesi – Handlirsch, 549, text-fig. 1885. Thus, after Article 21.6 of the Code of zoological 48 (new generic assignment). nomenclature (ICZN 1999), the date of publication to be 1919 Sthenaropoda minor – Handlirsch, 549, text-fig. adopted is 30 June 1885. Burnham (1983) indicated that 49 (assignment to a new species, new synonymy). Scudder’s (1885) work was published “early in April”, 1929 Archaeacridites bruesi – Vignon, 112, pl. 3, fig. 2. which supports its priority over Brongniart (1885). More- 1929 Sthenaropoda agnusi – Vignon, 113, pl. 3, fig. 2. over, Brongniart (1885: 68) noted that during the printing 1961 Sthenaropoda fischeri – Sharov, 245 (citation). of his work he received an exemplar of a new memoir 1968 Sthenaropoda fischeri – Sharov, 20, figs A, B from Scudder, dealing with new Paleozoic families and (restoration and original drawing). genera of Hexapoda. Judging from Scudder’s bibliogra- 1983 Gerarus fischeri – Burnham, 27–34, figs 14–17 phy, Brongniart very probably referred to the memoir of (revision; Gerarus fischeri, G. lerichei, and G. 1885 in which the genus Gerarus was published. agnusi synonymized). 1983 Gerarus bruesi – Burnham, 35–36, figs 18–19. Gerarus fischeri (Brongniart, 1885) 1992 Gerarus fischeri - Kukalová-Peck & Brauck- sensu Burnham (1983) mann, 2454, fig. 26 (original drawing). (Figs 1–7) 1992 Gerarus bruesi – Kukalová-Peck & Brauck- Synonymy list mann, 2454, fig. 27 (original drawing). 1992 Gerarus bruesi – Carpenter, 122, fig. 72.3a. 1885 Sthenaropoda fischeri – Brongniart, 59, pl. 1, 2002 Gerarus fischeri – Rasnitsyn, fig. 360 (photog- fig. 4 (short description with photography). raphy reproduced from Vignon 1929, drawings 1893 Oedischia fischeri – Brongniart, 437, pl. LI-35, after Burnham 1983). figs 5–6, pl. LII-36 fig 4. 2002b Gerarus bruesi – Béthoux & Nel, 35–37, figs 1906 Sthenaropoda fischeri – Handlirsch, 142, pl. XIV, figs 14–16. 13–14 (original drawing and photography). 1909a Archaeacridites bruesi – Meunier, 38–40, fig. 2 (original description). Material 1909b Sthenaropoda fischeri – Meunier, 43–44, figs Holotype: MNHN-LP-R.51128 (= IP 5), counter-imprint 3–4 (illustration of new material). R.51135 (= IP 7). Other specimens: R.51336 (= IP 23, 1909b Archaeacridites elegantissima – Meunier, 45–46, holotype of ‘G. lerichei’); R.51139 (= IP 19/21, holotype fig. 5 (original description; new synonymy). of ‘G. agnusi’); R.51146 (= IP 2); R.51164 (= IP 20; 1909c Archaeacridites elegantissima – Meunier, 140 holotype of ‘G. bruesi’); R.51227 (= IP 10); R.51242 (= (citation). IP 3); R.51269; R.51354 (= IP 4); R.11191. All at Labo- 1909d Archaeacridites bruesi – Meunier, 145, text-fig. ratoire de Paléontologie, Muséum National d’Histoire 16, pl. V, fig. 1 (new synonymy). Naturelle, Paris, France. Org. Divers. Evol. (2003) 3, 173–183 Wing venation of Gerarus fischeri (Panorthoptera) 175 Notes on synonymies fusion of RP with MA; RP with about 3–4 distal branch- Lameere (1917) designated the specimen MNHN-LP- es; M and its branches concave; position of origin of R.51139 as the holotype of G. agnusi after the figures of MA variable; origin of MP far distal of origin of CuA Meunier (1909d: pl. V, figs 4, 4a). After the same fig- from M + CuA; MP branched just distal of its origin, ei- ures, Handlirsch (1919) created the species G. minor. ther dichotomously branched or anteriorly pectinate; Surprisingly, Burnham (1983: 32) reports examining branches of MP usually distally branched, with different holotypes for these two species. Nevertheless, 3–8 branches reaching posterior wing margin; M + as Burnham synonymized G. agnusi with G. fischeri, we CuA, CuA and CuA + CuPaα convex; CuA + CuPaα also synonymize G. minor with G. fischeri. branched, either dichotomously or posteriorly pecti- We could not find the holotype of ‘G. elegantissima’ nate; branches of CuA + CuPaα usually distally in the collection of the MNHN. It is probably lost. But branched, with 5–9 branches reaching posterior wing the figure of Meunier (1909d: Pl. V, fig. 2) yields suffi- margin; CuP and branches of CuP concave, CuPaα con- cient information for the new synonymy with G. fischeri cave to convex; CuPaβ either well developed, or more (contra Burnham 1983): the long RP, the organization of or
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