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Org. Divers. Evol. 2, 313–318 (2002) © Urban & Fischer Verlag http://www.urbanfischer.de/journals/ode

Revision of the enigmatic Upper Campyloptera eatoni Brongniart, 1893 (Insecta: )

André Nel*, Aurélien Huguet

Muséum National d’Histoire Naturelle, Paris, France

Received 11 February 2002 · Accepted 27 August 2002

Abstract

Campyloptera eatoni Brongniart, 1893, the type species of the type of the Upper Carboniferous family Campylopteridae Handlirsch, is redescribed. It is not a as previously supposed, but an Odonatoptera with a specialized wing venation. Although it has a more position than the because of the absence of any nodal or subnodal structure, it has acquired a simple vein MA and a widening area between MP and CuA, convergently with the highly derived Discoidalia that includes the modern .A new diagno- sis is given for Campylopteridae and its type genus, Campyloptera Brongniart. Campylopterodea Rohdendorf, 1962 falls as a new junior syn- onym under Odonatoptera Martynov, 1932.

Key words: Insecta, Odonatoptera, Campylopteridae, Campyloptera eatoni Brongniart, phylogeny

Introduction Campyloptera is a genuine Odonatoptera with a highly specialized wing venation. Campyloptera eatoni Brongniart, 1893 is an enigmatic We follow the wing venation nomenclature of Riek insect from the Upper Carboniferous of Commentry, (1976), Riek & Kukalová-Peck (1984), amended by France. Originally considered as a Megasecoptera Kukalová-Peck (1991), Nel et al. (1993), and Bechly (Brongniart 1893), its systematic position was frequent- (1996). The higher classification of fossil and extant ly reassigned from this order to the Odonatoptera (under Odonatoptera is based on a criticism of the phylogenetic the name ‘Protodonata’) and back, as a ‘link’ between system of Bechly (1996). the two orders, or even as a different order Campy- lopterodea (Meunier 1907; Tillyard 1928; Carpenter 1931, 1943; Rohdendorf 1962). Most recently, Carpen- Order Odonatoptera Martynov, 1932 ter (1992) considered it as ‘Paleoptera order uncertain’. Campylopterodea Rohdendorf, 1962 syn. nov. Since the original study of Brongniart (1893), only Lameere (1917) and Carpenter (1943) revised the type Family Campylopteridae Handlirsch, 1906 specimen, but all of them failed to establish the exact Type genus: Campyloptera Brongniart, 1893. structure of its wing base. Bechly (pers. comm. 2002) New diagnosis: (1) wing narrow; (2) subpetiolate; (3) considered Campyloptera as an Odonatoptera: Odonato- ScP reaching costal margin nearly midway between base clada (= Lapeyriidae & Campylopteridae & Nodialata), and apex; (4) MA simple; (5) area between MP and CuA but after the wrong drawing of Carpenter (1943). With distally widened along posterior wing margin; (6) CuA the present study we can fill this gap, due to a careful distally zigzagged; (7) CuP distally simple and not and direct examination of this fossil under alcohol. zigzagged; (8) area between AA and posterior wing mar-

*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]

1439-6092/02/02/04-313 $ 15.00/0 Org. Divers. Evol. (2002) 2, 313Ð318 314 Nel & Huguet gin narrow; (9) MP and Cu basally separated but fused 11.7 mm wide; vein subcosta posterior (ScP) reaching well basal of respective points of separation of CuP and costal margin 33.8 mm from wing base, nearly midway of CuA from MP; (10) RP emerging from RP + MA well between base and apex; 11 cross-veins in ‘antenodal’ distal of base of arculus; (11) no nodal cross-vein (nodal area between costa (C) and ScP not aligned with the 11 Cr sensu Bechly 1996); (12) no subnodus. Character (6) cross-veins in area between radius anterior (RA) and is an autapomorphy. Characters (2), (4), (5) are apomor- ScP, except for the most basal one (‘primary antenodal phies convergently present in Discoidalia (see Discus- brace’); RA reaching costal margin 4.6 mm basal of sion below). wing apex; no pterostigma; no specialized nodal cross- vein and no subnodus opposite ‘nodus’ (i.e. point of fu- Genus Campyloptera Brongniart, 1893 sion between ScP and C) (Fig. 2); RP + MA separating Type species: Campyloptera eatoni Brongniart, 1893. from RA 8.0 mm from wing base (arculus); arculus very Diagnosis: See family diagnosis. oblique; radius posterior (RP) separating from media an- Campyloptera eatoni Brongniart, 1893 (Figures 1Ð3) terior (MA) well distant, 6.1 mm from base of arculus; Campyloptera eatoni Brongniart, 1893: 406, pl. 24, fig. RP with 2 concave posterior branches, RP3/4 and RP2, 3; in Megasecoptera. and 2 convex posterior branches, IR2 and IR1, these Campyloptera eatoni Brongniart Ð Handlirsch (1906: veins being rather regularly disposed; one row of cells 316, pl. 32, fig. 15); in Megasecoptera: Campylopteridae between IR1 and RP2 and between RP2 and IR2, but 5 nov. fam. Ð Meunier (1907: 521); in Protodonata. Ð rows of cells between IR2 and RP3/4 along posterior Lameere (1917: 163); in Megasecoptera. Ð Handlirsch wing margin; MA simple; only one row of cells between (1919: 580); in Megasecoptera: ‘? Fam. Campylopteri- MA and RP3/4; media posterior (MP) basally separated dae Handl.’. – Tillyard (1928: 161); in ‘Protodonata’: from cubitus (Cu) but these veins are fused 1.2 mm from Protagriidae Handlirsch, 1906, subfamily Campylopteri- wing base; MP + Cu with a strong anterior curve about nae. Ð Carpenter (1931: 102Ð103); in Megasecoptera, 5.4 mm distally; cubitus posterior (CuP) and cubitus an- after Lameere’s opinion. Ð Martynov (1932: 16); in fami- terior (CuA) separated from MP in this curve and reach- ly Protagriidae, with Protagrion Brongniart, 1894, as ‘in- ing analis anterior (AA), both looking like short, paral- termediate link between Megasecoptera and Meganeuri- lel, oblique cross-veins between AA and MP; CuP sepa- dae’. – Carpenter (1943: 548–552, fig. 4); in Megasec- rating again from AA 1.0 mm distal of their fusion; MP optera: Campylopteroidea nov. subordo. Ð Laurentiaux long and straight, more or less parallel to MA; CuA (1953: 445); in Megasecoptera: Campylopteroidea. Ð more or less parallel with MP but distally zigzagged; Rohdendorf (1962); in Campylopterodea nov. ordo. Ð CuP simple, concave and parallel to CuA and AA, with Carpenter (1992: 90); in Paleoptera, order uncertain. only one row of cells between it and AA; CuP reaching Material: Holotype specimen MNHN-LP-R.51236, posterior wing margin 37.7 mm from wing base; AA Laboratoire de Paléontologie, Muséum National d’His- reaching posterior wing margin 33.8 mm from wing toire Naturelle, Paris, France. base; anal area between AA and AP rather narrow, 2.6 Horizon and locality: Stephanian, Upper Carbonif- mm wide, with only one row of cells (2 near distal end of erous; Commentry, Allier, France. AA); AA and AP separated at wing base but basally par- Redescription (Figs 1Ð3): Several undescribed struc- allel; wing with a petiole looking more or less like that tures of the wing were visible under alcohol. Imprint and of Recent Zygoptera: ; width of petiole counterimprint of a single fore(?) wing 67.7 mm long, 5.1 mm; length of petiole 8.0 mm.

Fig. 1. Campyloptera eatoni, holotype specimen MNHN-LP-R.51236, general reconstruction of the wing. Scale bar represents 2 mm.

Org. Divers. Evol. (2002) 2, 313Ð318 Revision of Campyloptera 315

Discussion very close together’. Furthermore, in Campyloptera, these veins are fused to a long, double-barrel radial stem, Campyloptera has the synapomorphies of the as in Nodialata; ‘base of MA has lost its connection with Odonatoptera, i.e. ‘MP unbranched’; ‘subcostal vein the medial stem and is secondarily fused with RP’; ‘MP ScP fused with costal margin distincly basal of wing and Cu are at least shortly fused’; ‘the longitudinal wing apex’; ‘anal brace with a Z-like kink in the CuP (CuP- veins MP and CuA are not straight but undulating or crossing = ‘anal crossing’ sensu Fraser 1957) at the point even kinked’. of fusion with AA’(Bechly 1996). Therefore we propose Within this clade, Campyloptera would share with the to synonymize the order Campylopterodea Rohdendorf, ‘Eomeganisoptera’ Rohdendorf, 1962 (family 1962 with Odonatoptera. ‘Erasipteridae’ Carpenter, 1939) (sensu Bechly 1996) Within this clade, Bechly (1996) proposed to charac- the unique synapomorphy of this group proposed by this terise the Geroptera Brodsky, 1994 on the basis of the author, i.e. ‘ScP distinctly shortened, fusing with the following synapomorphies: (1) ‘archaedictyon reduced costal margin at a midwing position’, but this character and transformed into a regular polygonal meshwork of is homoplastic as it is also present in Eugeropteridae, crossveins’; (2) ‘ScP distinctly shortened, fusing with Paralogidae, and Panodialata. This ‘synapomorphy’ of the costal margin at a midwing position’. Bechly (1996) the ‘Eomeganisoptera’ is unknown in Erasipteroides noted that these characters are homoplastic, more pre- Brauckmann & Zessin, 1989 and Erasipterella Brauck- cisely, character (1) is convergent to Euodonatoptera and mann, 1983. Thus, the of the ‘Eomegan- character (2) is convergent to Erasipteridae, Paralogidae, isoptera’ is very poorly supported. This group is proba- and even more advanced . Thus this clade remains bly paraphyletic, as suggested by Bechly (1996) and our weakly supported. A potential different synapomorphy own recent studies (Hutin & Nel, in prep.). The of this clade could be the presence of 2 main branches of erasipterid Whalleyala bolsoveri (Whalley, 1979) AA (Hutin & Nel, in prep.). (Upper Carboniferous, Westphalian A, England) has a Nevertheless, Campyloptera has the synapomorphies subpetiolate narrow wing, very similar to that of Campy- of the Neodonatoptera Bechly, 1996 ( of loptera (Whalley 1979). Within this group, Erasipteron Geroptera), i.e. ‘RA and RP basally strictly parallel and larischi Pruvost, 1933 (type species), Erasipteroides valentini (Brauckmann, 1985), and Whalleyala Brauck- mann & Zessin, 1989 have a very short and distal fusion between MP and CuA, distal of separation of CuP from CuA (plesiomorphy) (Whalley 1979, Brauckmann 1983, Brauckmann et al. 1985, Brauckmann & Zessin 1989). This plesiomorphic character state is absent in Campy- loptera. It is also absent in Erasipterella piesbergensis Brauckmann, 1983 that has a basal fusion of MP with Cu. Thus, it may be not related to the other ‘Erasipteri- dae’. Erasipterella differs from Campyloptera in the Fig. 2. Campyloptera eatoni, holotype specimen MNHN-LP- presence of a distal part of AA that is strongly R.51236, ‘nodal region’. Scale bar represents 1 mm. zigzagged.

Fig. 3. Campyloptera eatoni, holotype specimen MNHN-LP-R.51236, photograph. Scale bar represents 1 cm.

Org. Divers. Evol. (2002) 2, 313Ð318 316 Nel & Huguet

The group Martynov, 1932 (sensu duced’ (after Bechly 1996). More precisely, the distal Bechly 1996) comprises the more derived Palaeozoic parts of these veins cannot be distinguished from the nu- ‘Protodonata’. This last author included into this group merous parallel posterior branches of CuA (after the the Namurotypidae, the Paralogidae, the Kargalotypi- original figures of Guthörl 1962). Kohlwaldia has a dae, the Kohlwaldiidae, and the Meganeuridae. Bechly broad cubito-anal area with numerous cross-veins. (1996) characterized this group on the basis of the fol- Campyloptera does not share these characters. Bechly lowing synapomorphies: (1) ‘large wing span of more (1996) characterized the Meganeuridae Handlirsch, than 200 mm (reversed in Paralogidae), correlated with 1906 as follows: (1) ‘presence of a characteristical an extremely increased number of cells in all wings (at oblique vein between RA and RP near the base of RP2’; least 500 cells)’; (2) ‘crowding of longitudinal veins (2) ‘increased number of more than 1.000 cells’ in the along the costal margin’; (3) ‘spines on longitudinal wings. Regarding (1), the oblique vein is in the same po- wing veins reduced (except along the costal margin)’; sition, nearly opposite the distal end of ScP, and thus (4) ‘very large body size’. Characters (1) and (4) are not could be homologous with the subnodus of the Panodi- shared by some undescribed Upper Meganeuri- alata (= Lapeyriiidae + Nodialata Bechly, 1996) (Nel et dae from the Lodève basin that have a wing span of less al. 1999). Thus, character (1) could in fact be a synapo- than 100 mm (Huguet & Nel in prep.). Character (3) is morphy of the Meganeuridae with the Panodialata. Re- unknown for numerous fossils. Character (2) is a quanti- garding (2), Bechly (1996) noted that this could be a tative one and should be better defined. The monophyly synapomorphy with Kargalotypidae and Kohlwaldiidae, of the Meganisoptera sensu Bechly (1996) remains reversed in Carpentertypinae. Furthermore, the small questionable (Huguet 2001). Upper Permian meganeurids do not have such a large Nel et al. (2001) transferred the Kargalotypidae number of cells. Campyloptera does not have the ‘mega- Zessin, 1983 in the Triadophlebiomorpha: Zygophlebii- neurid subnodal veinlet’. da. The Namurotypidae Bechly, 1996 ( sip- The Panodialata are characterized by the ‘presence of peli Brauckmann & Zessin, 1989) differs from Campy- a true odonatoid nodus, with a more or less oblique loptera in ‘ScP ending in costal margin distal of the mid- nodal Cr and subnodus’ (Nel et al. 1999). The Lapeyri- wing’ and in the ‘presence of a very short and distal fu- dae Nel et al., 1999 represents the most basal clade of sion of MP with Cu’ (plesiomorphies). Bechly (1996) the Panodialata. They have a vein MA with numerous characterized the Paralogidae Handlirsch, 1906 (two posterior branches (plesiomorphy), unlike the sister genera: Paralogus Scudder, 1893 and Oligotypus Car- group Nodialata Bechly, 1996 (Protanisoptera Carpen- penter, 1931) as follows: (1) ‘wings much shortened’; ter, 1931 and more advanced Odonatoptera) that have an (2) ‘costal margin concavely curved’; (3) ‘ScP distinctly unbranched vein MA (note that the Protanisoptera have shortened, fusing with the costal margin at a midwing a specialized vein IMA between MA and MP, Huguet et position’; (4) ‘first fork of RP very wide (RP1/2 and al. in press). Campyloptera has no nodal cross-vein RP3/4 strongly divergent)’; (5) ‘unique branching pat- (nodal Cr) and subnodus. Thus, it cannot be considered tern of CuA’; (6) ‘vestige of the median stem sup- as a Panodialata. But its MA is simple, suggesting possi- pressed’. Characters (1) and (2) are homoplastic. Char- ble relationships with the Nodialata. Thus, the two char- acter (4) is a quantitative one that would need to be rede- acters of Campyloptera, ‘absence of nodal structures’ fined. The branching pattern of CuA in Paralogus and ‘MA simple’, are in conflict. aeschnoides Scudder, 1893 is very particular, with a Campyloptera has another character, area between strong division of the main vein into 2 branches that de- MP and CuA distally widened near posterior wing mar- fine a distal triangular area with several secondary longi- gin, that is only present in the more advanced Discoidalia tudinal veins between them (Carpenter 1960). But the Bechly, 1996 (=Triadophlebiomorpha Pritykina, 1981 branching pattern of CuA in Oligotypus tillyardi Car- sensu Nel et al. 2001, Protozygoptera and Panodonata), penter, 1931 is very different, with only 3 distal branches characterized by a genuine discoidal cell structure with a of equal importance. Character (6) is not a strict synapo- vein MAb, but not in the Triadotypomorpha Bechly, morphy of these genera, as it is present in many more ad- 1996 (sensu Nel et al. 2001) (Triadotypidae Grauvogel & vanced Nodialata. The only character that could be a Laurentiaux, 1952), the Protanisoptera, the Lapeyriidae strict synapomorphy of the Paralogidae is the ‘costal and the Meganisoptera. It appears that the organization margin concavely curved’ just basal of apex of ScP. of the areas between MA, MP and CuA of Campyloptera Campyloptera only shares the characters (1) and (3) are autapomorphic and convergently similar to those of with this group, which are not sufficient to support a di- the Discoidalia. These structures of Campyloptera are rect relationship between Campylopteridae and Par- probably related to the narrowing and petiolation of its alogidae. The Kohlwaldiidae Guthörl, 1962 (one wing. Interestingly, the erasipterid Whalleyala also has a species: Kohlwaldia kuehni Guthörl, 1962) are charac- narrow wing (but less than in Campyloptera), and a terised by ‘the distal part of CuP and AA strongly re- broad area between MP and CuA and a relatively narrow

Org. Divers. Evol. (2002) 2, 313Ð318 Revision of Campyloptera 317 area between MA and MP (but less than in Campy- Brongniart, C. (1893): Recherches pour servir à l’histoire des loptera). The presence of a simple vein MA was so far insectes fossiles des temps primaires précédées d’une étude considered as a strict synapomorphy of the Nodialata. sur la nervation des ailes des insectes. Bull. Soc. Indust. The present study demonstrates that this character was at Minér. Saint-Etienne (3) 7: 491 pp. least subject to one convergence in Campyloptera. Carpenter, F. M. (1931): The Lower Permian of Kansas. Part 2. The orders , Protodonata, In conclusion, the structures of veins MA, MP and and Odonata. Am. J. Sci. (5) 21: 97Ð139. CuA alone are not sufficient for an attribution of Campy- Carpenter, F. M. (1943): Studies on Carboniferous insects loptera to the clades Panodialata or Nodialata. Its lack of from Commentry, France. Part 1. Introduction and families any oblique subnodal veinlet suggests that it could be in Protagriidae, Meganeuridae, and Campylopteridae. Bull. a more basal position than the Meganeuridae, if the Geol. Soc. Am. 54: 527Ð554. ‘meganeurid subnodal veinlet’ is considered homolo- Carpenter, F. M. (1960): Studies on North American Carbonif- gous to the subnodus of the Panodialata. Accepting this erous insects. 1. The Protodonata. Psyche 67: 98Ð109. hypothesis that is not clearly contradicted by anything, Carpenter, F. M. (1992): Superclass . Pp. xxii + 655 the Meganeuridae should be at least considered as the in: Moore, R. C. & Kaesler, R. L. (eds) Treatise on Inverte- sister group of the Panodialata and not included in the brate Paleontology, (R), Arthropoda 4, 3/4. Geological So- problematic ‘clade’ Meganisoptera sensu Bechly (1996) ciety of America and University of Kansas, Boulder. Fraser, F. C. (1957): A reclassification of the order Odonata, (Huguet & Nel in prep.). Because the position of based on some new interpretations of the venation of the Campyloptera is more basal than Bechly (pers. comm. wing. Hdbk R. Zool. Soc. New South Wales 12: 2002) supposed, the clade Odonatoclada Bechly, 2002 1Ð133. has to be synonymized with the Panodialata Nel et al., Guthörl, P. (1962): Zur Arthropoden-Fauna des Karbons und 1999. Campyloptera is in a more advanced position than Perms. 14. Kohlwaldia kühni n. g. n. sp. (Ins., Protodon.) the Erasipteridae and Namurotypidae, because of its aus dem Westfal D des Saarkarbons mit einer Zusammen- long basal fusion of MP with Cu, and in a more basal po- stellung der bis jetzt bekannten palaeozoischen sition than the Meganeuridae. But its relationships with Protodonatenarten. Annls Univ. Saraviensis Naturwiss.-Sci- the groups Paralogidae and Kohlwaldiidae remain unde- entia 10: 49-56. termined. Handlirsch, A. 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Meunier, F. (1907): Un Odonatoptère du Rhétien de Fort ous fossils from Argentina (Insecta: Odonatoidea) and basic Mouchard près des Arçures (Jura). Bull. Mus. Natl Hist. characters states in pterygote wings. Can. J. Zool. 62: Nat. Paris 13: 521Ð522. 1150Ð1166. Nel, A., Bethoux, O., Bechly, G., Martínez-Delclòs, X. & Pa- Rohdendorf, B. B. (1962): Podtip Mandibulata zhvalonosnye pier, F. (2001): The Permo- Odonatoptera of the chlenistonogie [Subphylum Mandibulata or Prototrachea- ‘protodonate’ grade (Insecta: Odonatoptera). Annls Soc. ta]. Pp. 17Ð374 in: Orlov, Yu. A. (ed.) Osnovy Paleontologii Entomol. Fr. 37: 501Ð525. [Fundamentals of Palaeontology]. Izdat. Akad. Nauk SSSR Nel, A., Gand, G. & Garric, J. (1999): A new family of Moscow 9: 560 pp. [English translation 1991. Smithsonian Odonatoptera from the continental Upper Permian: the Lapeyriidae (Lodève Basin, France). Geobios 32: 63–72. Institution Libraries and National Science Foundation, Nel, A., Martínez-Delclòs, X., Paicheler, J.-C. & Henrotay, M. Washington]. (1993): Les ‘Anisozygoptera’ fossiles. Phylogénie et classi- Tillyard, R. J. (1928): The evolution of the order Odonata. fication (Odonata). Martinia, Numéro Hors Série 3: 1–311. Recs Indian Mus. 30: 151Ð172. Riek, E. F. (1976): A new collection of insects from the Upper Whalley, P. E. S. (1979): New species of and Triassic of South Africa. Ann. Natal Mus. 22: 791Ð820. Protodonata (Insecta) from the Upper Carboniferous of Riek, E. F. & Kukalová-Peck, J. (1984): A new interpretation Britain, with a comment on the origin of wings. Bull. Brit. of Dragonfly wing venation based upon Early Carbonifer- Mus. Nat. Hist. (Geol.) 32: 85Ð90.

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