T H E F I R S T R E C O R D E D P R O T O Z Y G O P T E R A N I N S E C T S

F R O M T H E U P P E R P E R M I A N O F F R A N C E

by A N D R E N E L , G E O R G E S G A N D , J A C Q U E S G A R R I Cand J E A N L A P E Y R I E

ABSTRACT. The earliest known Odonatoptera: Protozygoptera from the Upper Permian of Lode`ve (France) aredescribed.Epilestes gallicasp. nov. belongs to the Permolestidae andLodevia longialatagen. et sp. nov. to thePermepallagidae. Both of these families were previously known from the Kazanian of Russia, suggesting a similar agefor the formation of Lode`ve.

WITHIN the Saxonian (Permian) of the Lode`ve Basin, few fossil-bearing strata have been reported (Feysand Greber 1972) (Text-figs 1–2). Fossil tracks of tetrapod vertebrates (Heyler and Lessertisseur 1963;Ellenberger 1983a, 1983b, 1984; Gand 1987, 1989) and of invertebrates (Debriette and Gand 1990; Gand1994), as well as sparse vertebrates and plants (Doubinger and Heyler 1959, 1975) have been found at thebase of the Saxonian, e.g. in the Rabejac Formation (sensuOdin 1986). Later, Ellenberger (1983a, 1983b)described a surface in the Salagou Formation (sensuOdin 1986) near La Lieude (at the top of theSaxonian) with many tetrapod vertebrate tracks and on which conchostracans, ‘limulids’ and ‘invertebratetracks (insects and worms)’ were also reported.

As there are invariably about 1000 m of virtually ‘azoic’ claystones and red siltstones between thefossiliferous strata of Rebejac and La Lieude, it was long thought, despite much prospecting, that theSaxonian gaps in the fossil record resulted from the aridity which reigned for much of Permian times. Arecent review of the substantial Lapeyrie collection now suggests that this assumption should be revised.

One of us (JL) and his family have collected a large number of crustacean prints (triopsids andconchostracans), bivalve shells, invertebrate tracks and insects from several supposed ‘azoic’ strata andthese go some way to filling the Saxonian gaps. These discoveries also enhance the fauna of this periodwhich was previously represented principally by ichnites (Ellenberger 1983a, 1983b, 1984; Gand 1987,1993). The insects are varied and include several new species of Blattoptera and Odonatoptera as well as afew other so far unreported taxa of family or order rank. The study of these fossils will enhance ourentomological knowledge of the Permian and enable us to fill in a few phylogenetic gaps, especially for theOdonatoptera.

This order was already highly diverse and with a long history by the Late Permian. Upper Permian andLower to Middle Triassic Odonatoptera belong to at least three suborders: Protanisoptera, Triadophle-biomorpha and Protozygoptera (Bechly 1995, p. 263). Whilst a true Protozygopteran has recently beendiscovered in the Westphalian of England (E. Jarzembowski, pers. comm.), the oldest known Odonata:Zygopteraþ Anisozygopteria (sensuBechly 1995) are represented in the Middle and Upper Triassic (Nelet al. 1993). Protozygoptera and Archizygoptera (¼ Kennedyomorphasensu Pritykina 1981) areconsidered as the sister group of modern Odonata (Bechly 1995). If, as according to Nel andHenrotay (1992), Archizygoptera (Protomyrmeleontidae) is a monophyletic group, well-definedby numerous venational synapomorphies, the monophyly of the Protozygoptera and their exactrelationships with Archizygoptera remain uncertain. The present study of new fossil wings of Proto-zygoptera from the Upper Permian of France adds some information for the phylogenetic study of thissuborder.

[Palaeontology, Vol. 42, Part 1, 1999, pp. 83–97, 1 pl.] q The Palaeontological Association

E N T O M O L O G I C A L S I T E S

The insect levels are specified in Text-figure 1 by the alphanumerics F23 (¼ Les Canals) and F31(¼ Bouisset). The fossil-bearing horizons belong to the second sedimentary cycle of the Permian series,classically termed ‘Saxonian’ in the literature (Kruseman 1962; Feys and Greber 1972; Laversanne1976; Alabouvetteet al. 1982; Odin 1986; Gand 1987; Chaˆteauneuf and Gand 1989). Sites F23 andF31 are located in the Octon facies (Kruseman 1962), part of the Salagou F5 Formation (Odin 1986)(Text-fig. 2A–B).

The Bouisset and Canals sites are some 100 m above level 500 (see Text-fig. 2B). The Permiancomprises two sedimentary cycles separated by the Rabejac conglomerate. Within the second cycle, the

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TEXT-FIG. 1. Location of the insect sites F23 (¼ Les Canals) and F31 (¼ Bouisset) in the southern and western parts ofthe Lodeve Permian basin.

TEXT-FIG. 2. A, the Lodeve Permian, after Odin (1986). Fo¼ formations; Ens¼ Members: LI¼ Lower Loiras, LS¼Upper Loiras, MA¼ Mas d’Alary, VI¼ Lower Viala, VS¼ Upper Viala; Cogema profile with V.A.¼ volcanic ashesin Roman numerals; En¼ environments.B, section of the ‘Saxonian’ with insect levels F23 Les Canals, withEpilestesgallica sp. nov., and F31¼ Bouisset, withLodevia longialatagen. et sp. nov. Odin¼ Odin 1986; R¼ bed-marks;a¼ insect wings, c¼ conchostracans, g¼ gastropods, i¼ Isopodichnussp., p¼ invertebrate tracks, t¼ burrows, ti¼

triopsids, v¼ footprints.C, F29 up to F20¼ new fossiliferous levels in the Saxonian.

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Salagou F5 Formation containing the insects is the thickest, attaining almost 2000 m in the south of thebasin. It is very monotonous and formed from multiple negative binary sequences of decimetric todecametric claystone and siltstone strata. Within the claystone small decimetric levels of siltstonessometimes occur, pinching out laterally. It is in these that one of us (JL) collected many insects,conchostracans and triopsids in association. On the basis of the presence of the crustaceans and from thesedimentology, these fossiliferous beds have been interpreted by Gandet al. (in pressa, in pressb) asrepresenting crescent-shaped megaripples in temporary water bodies belonging to a semi-arid playa orpediplain environment. The Arie`ges site (¼ F19), as an example, comprises two lenses of very hard, redsiltstones, laminated with carbonate cement and separated vertically by some ten metres. Each lensterminates upwards with a level of desiccation cracks acting as markers in the formation. That of lens I canbe traced for a hundred metres but that of lens II is lost in the claystones after some ten metres of lateralextension. The dimensions of I and II are respectively 4 m and 6 m length by 0·5 m and 0·3 m maximumthickness. Text-figure 3 shows a cross section through lens I. Several sequences of poorly sorted siltstonescan be recognized, overlain by beds containing triopsids, conchostracans and undeformed insects.

The suggested ages of these sites have varied as discoveries have been made and cross-referencesconsidered. Based on various criteria the following ages have been suggested for the Salagou Formation,which contains insect-bearing levels.1. Saxonian–Thuringian, on tectonic and petrographic criteria. An unconformity followed by a conglom-erate allowed the Permian to be subdivided into Autunian and Saxonian–Thuringian (Feys and Greber1972).2. Upper Saxonian, from the distribution of tetrapod vertebrate traces (Gand 1987), after Haubold(1971).3. Probably post-Zechstein 1, as the insect beds overlie microflora levels ascribed to the Zechstein 1 byDoubingeret al. (1987).4. Perhaps post-Leonardian and therefore post-Kungurian, after the correlation chart of Ya Leven (1992),as the Rabejac Formation, underlying the Salagou Formation (Text-fig. 2A), has yieldedSupaiasp. whichdoes not survive the Leonardian (Doubinger and Heyler 1975).5. Thuringian, from recent palaeomagnetic data (Diego-Orozco 1994).6. The present discoveries of two Odonatoptera which are close to taxa firmly placed in the RussianPermian stratigraphy, make it possible to suggest the age of these beds at Arie`ges, Canals and Bouisset aslowermost Kazanian.

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TEXT-FIG. 3. Fossiliferous site F19; perspective view of lens I due to the filling of a gutter; j¼ bedding-plane and l¼fossiliferous beds.

S Y S T E M A T I C P A L A E O N T O L O G Y

SuborderPROTOZYGOPTERATillyard, 1925

Type family.Kennedyidae Tillyard, 1925.

Phylogenetic position.Pritykina (1981, p. 34) considered that Protozygoptera and ArchizygopteraHandlirsch, 1906 (Promomyrmeleontidae) should be put together in the infraorder KennedyomorphaPritykina, 1981. Carpenter (1992) concurred but renamed this group Archizygoptera. Nel and Henrotay(1992) considered that Protozygoptera requires revision and is probably not directly related to theProtomyrmeleontidae. Thus, these authors restricted Archizygoptera to the sole family Protomyrmeleon-tidae. Bechly (1995) also considered that the relative phylogenetic positions of ‘true Odonata’,Protozygoptera and Archizygoptera remain unresolved. Bechly (1996), however, proposed a newphylogenetic classification of these taxa, including Archizygoptera in Protozygoptera as sister-group ofPermagrionoidea, Protozygoptera being the sister-group of Panodonata, viz. more Recent Odonatoptera.

Family PERMOLESTIDAEMartynov, 1932

Type genus. PermolestesMartynov, 1932 (type speciesPermolestes gracilisMartynov, 1932; Kazanian, UpperPermian, Arkhangelsk District, Siberia).

Other genera. EpilestesMartynov, 1937 (type speciesEpilestes kargalensisMartynov, 1937; lowermost Kazanian,Permian, Kargala, Semenovski, Siberia),ScytolestesMartynov, 1937 (type speciesScytolestes stigmalisMartynov,1937; Permian, Kargala, Semenovski, Siberia) andSolikamptilon Zalessky, 1948 (type speciesSolikamptilonremuliformeZalessky, 1948; Permian, Sibera). Carpenter (1992) included the genusSushkiniaMartynov, 1930(type speciesS. parvulaMartynov, 1930 andS. elongataMartynov, 1930 from the Ufimian Horizon, Kungurian,Permian, Tikhie Gory, Kama River, Kazan, Russia) which was originally considered by Martynov (1930) asKennedyidae. The attribution ofSushkiniato Permolestidae is uncertain because only the distal parts of the wingsof both S. parvulaand S. elongataare known, and so the wing venational synapomorphies of Permagrionoidea(¼ Permolestidaeþ Permagrionidae) are unknown in these taxa. Furthermore,S. parvulaandS. elongatashare withKennedyidae two synapomorphies, viz. no more than one crossvein beneath the pterostigma, and few postnodals.Thus, our present state of knowledge strongly suggests thatSushkiniais more closely related to Kennedyidae than toPermolestidae.

GenusEPILESTESMartynov, 1937

Type species. Epilestes kargalensisMartynov, 1937.

Epilestes gallicasp. nov.

Plate 1, figure 2; Text-figure 4

Derivation of name.After the ancient name of France.

Holotype.Specimen Ld LAP 110, coll. Dr Lapeyrie, Muse´e Fleury, Lode`ve.

Type locality and horizon.Upper Permian, Kazanian, Les Canals (¼ F23), Lodevois, France.

Diagnosis.The fore- (?) wing of this species is characterized as follows: long but rather wide, wellpetiolated; cubital area broad, with three rows of cells between CuA and posterior wing margin; CuA withnumerous long branches; main veins are long and straight but without intercalary longitudinal veins

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between them; MP separated from Cu at wing base; pterostigma long and narrow; wing apex acute; ninepostnodal and three postsubnodal crossveins; oblique vein m-cu opposite MAb.

Description.Counterpart of a complete wing, 29·0 mm long, 5·2 mm wide. Convex veins appear concave and vice-versa. No trace of coloration preserved. Distance from base to nodus 11·2 mm, from nodus to apex 17·7 mm. Nodus in abasal position. Distance from base to arculus 4·6 mm. Petiolus long and narrow, 5·3 mm long and 1·3 mm wide.MPþ Cu and AAþ AP strongly approximate in petiolus. CuP not preserved. MP basally separated from Cu, closelyparallel and distally fused with it, 2·6 mm from wing base. At wing base, MP weakly concave and Cu strongly convex.RP separated from MA 0·5 mm distal of base of arculus, strongly oblique. First crossvein between MA and MP 0·8 mmlong and strongly convex, looking like a posterior branch MAb of MA (Nelet al. 1993) distally closing an anteriorlyopen discoidal space (Text-fig. 4). CuA and MP separate 0·5 mm distal of end of petiolus and distinctly basal of MAb,making a sharp angle. CuA strongly convex, with numerous posterior branches and concave short veins between them.Cubital area rather broad, 1·5 mm wide, with three rows of cells between CuA and posterior margin. Main veins, i.e.MP, MA, RP0 0, IR2, RP2, and IR1 simple and nearly straight. CuA reaching posterior margin at 77 per cent., MP at 85per cent. and MA at 89 per cent. of wing length. RP0 0 beginning 7·1 mm distal of arculus and 0·5 mm distal ofsubnodus, IR2 1·9 mm distal of base of RP0 0, RP2 4·2 mm distal of that of IR2, and IR1 2·7 mm distal of that of RP2.IR1 7·5 mm long. A single row of cells in areas between main longitudinal veins CuA, MP, MA, RP0 0, IR2, RP2, IR1and RP1. Wing apex acute. No oblique crossvein ‘O’ between IR2 and RP2. Although nodus poorly preserved, nodalcrossvein Cr between ScP and RA very clear and subnodus Sn distinctly oblique. Pterostigma poorly preserved butlong and narrow,c. 3 mm long. No oblique pterostigmal brace. Postnodal area poorly preserved, but with numerouspostnodal crossveins, not aligned with postsubnodal crossveins between RA and RP1. Antenodal area poorlypreserved, but two primary antenodal crossveins AX1 and AX2 visible, 2·0 mm apart. AX2 0·5 mm basal of arculus.If present, secondary antenodal crossveins not clearly preserved. Only one crossvein in area between RA and RP, basalof subnodus. AA completely fused with AP. Opposite MAb, A strongly concave and very short oblique crossvein m-cu(sensuBechly 1996), 0·1 mm long, between MP and CuA, and looking like a posterior branch of MP rather than otherconvex crossveins between MP and CuA (Text-fig. 4).

Remarks.In comparison with other Permolestidae, this new fossil taxon is very similar to the wings of theProtozygoptera: PermolestidaeEpilestes, Scytolestesand Permolestes. The differences between thesegenera andEpilestes gallicasp. nov. are as follows. There is no secondary longitudinal vein between all

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TEXT-FIG. 4. Epilestes gallicasp. nov.; holotype, Ld LAP 110, Fleury Museum in Lode`ve; wing venation. Scale barsrepresent 2 mm.

main veins, unlikePermolestes graciliswhich has one between RP2 and IR2 and between RP00 and MA.MA of P. gracilis is distally zigzagged and indistinct in the area between RP00 and MP. IR1 is straight andRP2 is not secondarly branched on IR2, unlikeScytolestes stigmalis. The wing ofE. gallica sp. nov. issomewhat smaller (29 mm long) than that ofE. kargalensis(31 mm long); there are three rows of cells inthe cubital area instead of two inE. kargalensis; the wing apex is more acute than inE. kargalensis; andthere are fewer subpostnodal crossveins. All other venational characters are identical in the two species.

Martynov (1932) noted that the hindwing pterostigma ofP. gracilis is distinctly shorter and in a moredistal position than that of the forewing. From this remark, it is possible to infer that the holotype specimenof E. gallica might be a forewing.

Phylogenetic affinities within Protozygoptera.Bechly (1996) considered Protozygoptera to be the sistergroup of Panodonata (¼ Tarsophlebioideaþ [Zygopteraþ Anisozygopteria]): the sister group of(Protozygopteraþ Panodonata¼ Stigmoptera) being the Triadophlebiomorpha. Bechly’s two mainsynapomorphies of Stigmoptera, absent in Triadophlebiomorpha, are: (1) ‘the distal free part of veinCuP is secondarily absent’ (suppressed or fused with the anal vein) and (2) ‘a sclerotized pterostigma ispresent’. These synapomorphies are present inE. gallica. Bechly (1996) proposed the followingsynapomorphies for the Protozygoptera (¼ [PermagrionidaeþPermolestidae]þ Archizygoptera): (1)‘strongly petiolated wings (convergent to fossil Triadophlebiomorpha and many Zygoptera) with a verylong petiolus’; (2) ‘nodus shifted in a more basal position (than in Triadophlebiomorpha)’; (3) ‘regressivedevelopment of the nodus’; (4) ‘number of antenodals reduced to 2–3 that are more or less aligned’; (5)‘CuP-crossing shifted in a very basal position’. All known Permolestidae (includingE. gallica) sharecharacters (1) to (4) with other Protozygoptera but character (5) is figured only inP. gracilis. Martynov(1937) and Zalessky (1948) did not mention this vein inE. kargalensis, S. stigmalisandS. remuliforme.Character (5) is also unknown inE. gallica sp. nov. Furthermore, the new species shares withProtozygoptera the separation between CuA and MP distinctly basal of MAb (plesiomorphy).

Within Protozygoptera, Permagrionoidea (¼ Permagrionidaeþ Permolestidae) are characterized by thesynapomorphy, shared byE. gallica, of nearly aligned arculus and AX2 (convergently acquired by someZygoptera). Archizygoptera (sister group of Permagrionoidea within Protozygoptera) is characterized bythe nodal and subnodal crossveins being transverse, non-aligned and reduced. This reduction of the nodalstructures is absent inE. gallica. Following Bechly (1996), the Permagrionidae (including the UpperPermian type genus and speciesPermagrion falklandicusTillyard, 1928; Bodie Creek Beds, Gondwana,Falkland Islands) are characterized by the synapomorphies: (1) ‘apex of the wing strongly falcate’; (2)‘basal and distal margin of the pterostigma strongly oblique’; (3) ‘basal vestige of medial stem Msuppressed’; (4) ‘concave oblique crossvein m-cu aligned with MAb as a pseudo-subdiscoidal veinlet’.

Bechly (1996) recognized no wing venational synapomorphy of the Permolestidae. In the followingdiscussion, we list their possible synapomorphies.1. The polarity of character (2) (pterostigmal structures) is uncertain, because the pterostigma is absent inTriadophlebiomorpha, sister group of Stigmoptera. The very long pterostigma of Permolestidae, with itsnon-oblique basal margin and its distal margin distinctly oblique, is a very peculiar structure, also presentin Kennedyidae and Bakteniidae (Archizygoptera; Bechly 1996). The pterostigmal structure of Perma-grionidae (basal and distal margins oblique and parallel) is shared by Archizygoptera: Protomyrmeleonti-dae and Panadonata. If it is supposed that the pterostigmal structure of Permagrionidae is an apomorphy, itwould imply that it has been acquired convergently by Protomyrmeleontoidea and Panodonata. On theother hand, if the pterostigmal structure of Permolestidae is an apomorphy, it would also imply twoconvergences with Kennedyidae and Bakteniidae. Thus, it is still impossible to decide which of the twosolutions is the more parsimonious.2. The base of CuA is nearly opposite the arculus inPermagrionTillyard, 1928, unlike in Permolestidaeand Epilestes gallicasp. nov. However, as a situation similar to that in Permolestidae occurs inTriadophlebiidae (Triadophlebiomorpha) and Kennedyidae, it is probably a symplesiomorphy ofPermolestidae.3. Veinlet m-cu is aligned with MAb inPermagrionandE. gallicaand distal of MAb inPermolestes. Thisstructure was not mentioned by Martynov (1937) and Zalessky (1948) for other Permolestidae

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(E. kargalensis, S. stigmalisandS. remuliforme). Thus, it is very difficult to determine the exact value ofthis character. Furthermore, the triadophlebioideanTriadophlebia madygenicaPritykina, 1981 shares withPermagrionthe alignment of m-cu with MAb, suggesting that it is a plesiomorphy.4. All Permolestidae, except perhapsE. kargalensis, have distinctly fewer postsubnodal than postnodalcrossveins (three against five forP. gracilis; three against six forS. stigmalis; three against nine forE. gallica). The same situation occurs in Kennedyidae. The postnodal and subpostnodal areas ofE. kargalensisseem to be poorly preserved, as several postnodal crossveins that should be present are notfigured and several postsubnodals are indicated as dotted lines by Martynov (1937). There are as manypostsubnodal crossveins as postnodal crossveins inPermagrion. The same situation occurs in Triadoph-lebioidea and Protomyrmeleontoidea (¼ Bakteniidae and Protomyrmeleontidae) and Panodonata. Thus, itis more likely that the reduction of the number of postsubnodal crossveins is a synapomorphy ofPermolestidae, which is also present in Kennedyidae (synapomorphy or convergence?).

As E. gallicahas none of the ‘strong’ apomorphies ofPermagrion(characters (1) and (3) listed above)and as it shares one possible synapomorphy of the Permolestidae, we propose to include it in this family. Itwill be possible to verify this hypothesis after revision of the holotype ofEpilestes kargalensis.

Family PERMEPALLAGIDAE Martynov, 1938

Type genus. PermepallageMartynov, 1938 (type speciesPermepallage angustissimaMartynov, 1938, from thePermian, Kazanian, Sojana River, Arkhangelsk district, Russia).

Other species.Rohdendorfet al. (1961, p. 89, text-fig. 37) described a ‘Permepallagidea fam., gen., sp.’ from thePermian of Russia, which is anincertae sedisbecause it is based on a poorly preserved costal portion of a wing.

Wing venational autapomorphies(modified from Bechly 1996). Wings extremely slender; secondaryantenodals present (possible reversal ?); very long and concave oblique veinlet m-cu; postnodal andpostsubnodal crossveins very numerous.

GenusLODEVIA gen. nov.

Derivation of name.From the town Lode`ve (Herault, France).

Types species. Lodevia longialatasp. nov.

Diagnosis. The wing venation ofLodeviagen. nov. is similar to that ofPermepallage, differing in: onlyone concave basal crossvein between MP and CuA, instead of two; absence of intercalary veins betweenbranches of RP; obliquity of the subnodus, instead of being nearly perpendicular to RA as inPermepallage; obliquity of MAb directed towards wing apex, instead of being directed towards base asin Permepallage. The presence of numerous antenodal crossveins, considered by Bechly (1996) as anapomorphy ofPermepallage, is unknown inLodevia.

Lodevia longialatasp. nov.

Plate 1, figure 1; Text-figures 5–6

Derivation of name.After its very long and slender wing.

Material. Holotype specimen Ld LAP 111, paratype specimen Ld LAP 424, coll. Dr Lapeyrie, Muse´e Fleury, Lode`ve.

Type locality and horizon.Upper Permian, Kazanian, holotype from Bouisset (¼ F31), paratype from Le Moural D(¼ F21D), Lodevois, France.

Diagnosis.As for genus (this is the only recognized species).

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DescriptionHolotype (Text-fig. 5). Impression of a nearly complete wing very long and slender, about 38 mm long and 4·9 mmwide, which is only broken opposite the nodus. No trace of coloration preserved. Distance from base to nodus about15 mm, from nodus to apex 23 mm. Nodus in a basal position. Distance from base to arculus 6·8 mm. Petiolus long andvery narrow, 6·5 mm long and 1·0 mm wide. MPþ Cu and AAþ AP long parallel but not strongly approximate inpetiolus. Transverse CuP not visible due to anal area covered by sediment, but CuP with no distal free part. MP notbasally separated from Cu. MPþ Cu strongly concave. RP separated from MA by a long distance, 0·7 mm distal ofbase of the strongly oblique arculus. First crossvein MAb between MA and MP strongly convex and distally closing abasally open discoidal space. MA and MP strongly approximate and diverging again distally, thus seeming to close thediscoidal space. CuA and MP separated 0·4 mm distal of base of arculus, making a sharp angle. CuA strongly convex,with numerous posterior short branches but without concave veins between them. Cubital area narrow, 0·8 mm wide.Main veins, i.e. CuA, MP, MA, RP0 0, IR2, RP2 and IR1 all nearly straight. CuA reaching posterior margin at 70 percent., MP at 84 per cent. and MA at 90 per cent. of wing length. RP0 0 probably beginning slightly distal of nodus. IR2beginning about 4 mm distal of base of RP0 0, RP2 6·2 mm distal of that of IR2. and IR1 beginning 4·0 mm distal of thatof RP2. IR1 9·7 mm long. One row of cells in the areas between the main veins, i.e. posterior margin, CuA, MP, MA,RP0 0, IR2, RP2, IR1 and RP1. Wing apex rounded. One oblique crossvein between IR2 and RP2, five cells distal ofbase of RP2. Nodus not preserved. Pterostigma long and narrow, 3·7 mm long, covering five cells, with its proximalside making a right angle with RA and its distal side oblique. No oblique pterostigmal brace. Postnodal area well-preserved with 14 postnodal crossveins, not aligned with the 12 postsubnodal crossveins between RA and RP1.Antenodal area poorly preserved, covered by sediment; thus the two primary antenodal crossveins not visible. Novisible crossvein in area between RA and RP, basal of subnodus. AA completely fused with AP. A strongly concaveand oblique vein m-cu between MP and CuA, rather long, 0·7 mm long and nearly opposite MA2.

Paratype (Text-fig. 6). Impression of a costal part of a wing very long and slender. No trace of coloration preserved.Main veins, i.e. CuA, MP, MA, RP0 0, IR2, RP2 and IR1, all nearly straight with one row of cells in the areas betweenthem. Nodus well preserved with the subnodus distinctly oblique and not aligned with nodal crossvein Cr. Postnodalarea well-preserved with 16 visible postnodal crossveins, not aligned with the 15 postsubnodal crossveins between RAand RP1. Two visible crossveins in area RA and RP, basal of subnodus. RP0 0 begins one cell distal of the subnodus.

Remarks.We attribute these two specimens to the same species because they share the same organizationof the main veins in the median part of the wing and the absence of any intercalary veins between them.Lodevia longialatagen. et sp. nov. shares nearly all the venational synapomorphies of the Stigmoptera:Protozygoptera, viz. no distal free part of CuP, a sclerotized pterostigma between C and RA, wing stronglypetiolated with AA and AP completely fused, petiolus very long, nodus shifted in a basal position. Itsposition within Protozygoptera is more difficult to establish. Nevertheless, it can be provisionally related tothe Permepallagidae because of the unique synapomorphies within Protozygoptera: principally, numerouspostnodal and subpostnodal crossveins, a very long and slender wing and oblique m-cu very long(distinctly longer than MAb).Lodeviagen. nov. andPermepallagealso share the presence of numerouscrossveins beneath the pterostigma between RA and RP1 (probable symplesiomorphy), the base of RP00

one cell distal of the subnodus and the presence of numerous crossveins between RA and RP basal of thesubnodus. Nevertheless, this attribution is still open because the family characters of Permepallagidae arenot at all clear.

Martynov (1938, p. 50) considered the presence of a basal enigmatic longitudinal concave (?) vein in thepetiolus, fused with the anal margin about midway of the petiolus, and that of a small crossvein, betweenMPþ Cu and the previous enigmatic vein, as the main diagnostic character ofPermepallage. As it is

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TEXT-FIG. 5. Lodevia longialatagen. et sp. nov.; holotype, Ld LAP 111, Fleury Museum in Lode`ve; wing venation.Scale bar represents 2 mm.

possible to homologize the small crossvein as CuP, because of its position, then the basal enigmaticlongitudinal concave (?) vein would be AA basally separated from AP and distally fused again with it. But,AA is a convex vein in all other Odonatoptera. Thus, we think that the convexity of this enigmatic vein wasmisinterpreted by Martynov, or it is only an artefact. Re-examination of the holotype ofPermepallageangustissimawill be necessary to solve this problem. Nevertheless, it is not possible to definePermepallagidae using this dubious character, which is, anyway, poorly preserved in the holotype ofLodevia longialatagen. et sp. nov.

Besides, the presence of intercalary longitudinal veins between the branches of RP, which is consideredby Bechly (1996) as an apomorphy ofPermepallage, is not sufficient to excludeLodevia fromPermelpallagidae on account of similarities in Recent Zygoptera: Coenagrionidae. In them, the numberof intercalary veins varies greatly through the different genera so that this character is of little significanceat family level. Thus, includingLodeviain a new family is not necessary, and also because it would add noinformation to the relationships between it andPermepallage.

C O N C L U S I O N S

Previous to the description ofEpilestes gallicasp. nov. andLodevia longialatagen. et sp. nov.,Arctotypusverneti Laurentiaux-Vieira and Laurentiaux, 1963 (Meganeuridae Tupinae, Upper Permian, AlpesMaritimes, France), was the only described odonatopteran from the Upper Permian of France. Asundescribed Tupinae are also present in the Lode`ve’s Formation, together with a new undescribedfamily, related to the Protanisoptera, the odonatan fauna from the Saxonian of France appears to have beenabundant and diverse (Gandet al. in pressb; Nel et al. in press).

The two newly described taxa are attributed to the families Permepallagidae and Permolestidae whichhave been, until now, described only from the Upper Permian (Kazanian) of Russia. Thus, it is possible toinfer tentatively a similar age for a part of the Upper Salagou Formation where they are found.Nevertheless, the present fossil record of the families of Protozygoptera is not congruent (sensuHuelsenbeck1994) with the phylogeny proposed by Bechly (1996) (see Table 1 and Text-fig. 7). Their sister-grouprelationships with the Archizygoptera imply that Permagrionoidea were present during the Early Permian.Thus, the presence of Permolestidae in the Upper Permian has less stratigraphical importance than might besupposed. The inferred Kazanian age needs further evidence based on the study of the other insects and theircomparison with the entomofaunas of the Permian of Russia and North America.

92 P A L A E O N T O L O G Y , V O L U M E 4 2

E X P L A N A T I O N O F P L A T E 1

Fig. 1. Lodevia longialatagen. et sp. nov.; holotype, Ld LAP 111, Fleury Museum in Lode`ve; Site F31¼ Bouisset,Salagou Formation;×3·5.

Fig. 2.Epilestes gallicasp. nov.; holotype, Ld LAP 110, Fleury Museum in Lode`ve; Site F23¼ les Canals, SalagouFormation;×4·25.

TEXT-FIG. 6. Lodevia longialatagen. et sp. nov.; paratype, Ld LAP 424, Fleury Museum in Lode`ve; wing venation.Scale bar represents 2 mm.

P L A T E 1

NEL et al., Lodevia, Epilestes

94 P A L A E O N T O L O G Y , V O L U M E 4 2

TABLE 1. Permo-Triassic taxa attributable to Protozygoptera.

PERMAGRIONOIDEAPERMOLESTIDAE Martynov, 1932

PermolestesMartynov, 1932Permolestes gracilisMartynov, 1932 (Upper Permian)

EpilestesMartynov, 1937Epilestes kargalensisMartynov, 1937 (Upper Permian)Epilestes gallicasp. nov. (Upper Permian)

ScytolestesMartynov, 1937Scytolestes stigmalisMartynov, 1937 (Upper Permian)

SolikamptilonZalessky, 1948Solikamptilon remuliformeZalessky, 1948 (Permian)

PERMAGRIONIDAE Tillyard, 1928PermagrionTillyard, 1928

Permagrion falklandicusTillyard, 1928 (Upper Permian)

ARCHIZYGOPTERAKENNEDYIDAE Tillyard, 1925

KennedyaTillyard, 1925Kennedya mirabilisTillyard, 1925 (Carpenter 1931, 1933, 1939, 1947) (Lower Permian)Kennedya tillyardiCarpenter, 1939 (Permian)Kennedya reductaCarpenter, 1939 (Permian)Kennedya fraseriCarpenter, 1947 (Permian)Kennedya carpenteriPritykina, 1981 (Triassic)Kennedya gracilisPritykina, 1981 (Triassic)Kennedyasp. (Tasch and Zimmerman 1959, 1962) (Permian)

ProgoneuraCarpenter, 1931Progoneura minutaCarpenter, 1931 (Permian)Progoneura nobilisCarpenter, 1947 (Permian)Progoneura venulaCarpenter, 1947 (Permian)Progoneurasp. (Carpenter, 1947) (Permian)

SuschkiniaMartynov, 1930Suschkinia parvulaMartynov, 1930 (Permian)Suschkinia elongataMartynov, 1930 (Permian)

OpterSellards, 1909Opter brongniartiSellards, 1909 (Permian)

PERMEPALLAGIDAE Martynov, 1938PermepallageMartynov, 1938

Permepallage angustissimaMartynov, 1938 (Upper Permian)Lodeviagen. nov.

Lodevia longialatasp. nov. (Upper Permian)Permepallagidea fam., gen., sp. (Rohdendorfet al. 1961) (Upper Permian)

PROTOMYRMELEONTOIDEABATKENIIDAE Pritykina, 1981

BatkeniaPritykina, 1981Batkenia pusillaPritykina, 1981 (Triassic)

TerskejaPritykina, 1981(ProtomyrmeleontidaesensuPritykina 1981; uncertain position after Nel and Henrotay 1992; in Batkeniidae after Bechly1996)

Terskeja paulaPritykina, 1981 (Triassic)Terskeja pumilioPritykina, 1981 (Triassic)Terskeja tenuisPritykina, 1981 (Triassic)

VoltzialestesNel, Papier, Grauvogel-Stamm and Gall, 1997Voltzialestes triasicusNel, Papier, Grauvogel-Stamm and Gall, 1997 (Lower Triassic)

PROTOMYRMELEONTIDAE Handlirsch, 1906TillyardomyrmeleonHenrotay, Nel and Jarzembowski, 1997

Tillyardomyrmeleon petermilleriHenrotay, Nel and Jarzembowski, 1997 (Middle Triassic)TriassagrionTillyard, 1922

Triassagrion australienseTillyard, 1922 (Upper Triassic)

Acknowledgements.This work is a contribution to the team ‘Pale´ontologie et Biostratigraphie du Permien’ of theUMR/CNRS ‘Paleontologie analytique et Ge´ologie sedimentaire’. It is supported by the UMR 5561/CNRS. Our greatthanks to Mr Godon and Mme Bussie`re of the Centre des Sciences de la Terre de l’Universite´ de Bourgogne, for thephotographs of the fossils and the preparation of the figures.

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ANDRE NEL

Laboratoire d’EntomologieMuseum national d’Histoire naturelle

45 rue de Buffon, F-75005, Paris, France

GEORGES GAND

UMR 5561 du CNRSPaleontologie analytique etGeodynamique se´dimentaire

Universitede BourgogneCentre des Sciences de la Terre

6, boulevard Gabriel. F-21000, Dijon, France

JACQUES GARRIC

16 rue des Azale´es; La ChamberteF-34070, Montpellier, France

JEAN LAPEYRIE

Corniche de FontbonneF-34700, Lode`ve, France

Typescript received 21 March 1997Revised typescript received 20 April 1998

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