THE FIRST RECORDED PROTOZYGOPTERAN INSECTS FROM THE UPPER PERMIAN OF FRANCE

Transcription

1 THE FIRST RECORDED PROTOZYGOPTERAN INSECTS FROM THE UPPER PERMIAN OF FRANCE by ANDRÉ NEL, GEORGES GAND, JACQUES GARRIC and JEAN LAPEYRIE ABSTRACT. The earliest known Odonatoptera: Protozygoptera from the Upper Permian of Lodève (France) are described. Epilestes gallica sp. nov. belongs to the Permolestidae and Lodevia longialata gen. et sp. nov. to the Permepallagidae. Both of these families were previously known from the Kazanian of Russia, suggesting a similar age for the formation of Lodève. WITHIN the Saxonian (Permian) of the Lodève Basin, few fossil-bearing strata have been reported (Feys and 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; Gand 1994), as well as sparse vertebrates and plants (Doubinger and Heyler 1959, 1975) have been found at the base of the Saxonian, e.g. in the Rabejac Formation (sensu Odin 1986). Later, Ellenberger (1983a, 1983b) described a surface in the Salagou Formation (sensu Odin 1986) near La Lieude (at the top of the Saxonian) with many tetrapod vertebrate tracks and on which conchostracans, limulids and invertebrate tracks (insects and worms) were also reported. As there are invariably about 1000 m of virtually azoic claystones and red siltstones between the fossiliferous strata of Rebejac and La Lieude, it was long thought, despite much prospecting, that the Saxonian gaps in the fossil record resulted from the aridity which reigned for much of Permian times. A recent 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 and conchostracans), bivalve shells, invertebrate tracks and insects from several supposed azoic strata and these go some way to filling the Saxonian gaps. These discoveries also enhance the fauna of this period which 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 a few other so far unreported taxa of family or order rank. The study of these fossils will enhance our entomological knowledge of the Permian and enable us to fill in a few phylogenetic gaps, especially for the Odonatoptera. This order was already highly diverse and with a long history by the Late Permian. Upper Permian and Lower to Middle Triassic Odonatoptera belong to at least three suborders: Protanisoptera, Triadophlebiomorpha and Protozygoptera (Bechly 1995, p. 263). Whilst a true Protozygopteran has recently been discovered in the Westphalian of England (E. Jarzembowski, pers. comm.), the oldest known Odonata: Zygoptera þ Anisozygopteria (sensu Bechly 1995) are represented in the Middle and Upper Triassic (Nel et al. 1993). Protozygoptera and Archizygoptera (¼ Kennedyomorpha sensu Pritykina 1981) are considered as the sister group of modern Odonata (Bechly 1995). If, as according to Nel and Henrotay (1992), Archizygoptera (Protomyrmeleontidae) is a monophyletic group, well-defined by numerous venational synapomorphies, the monophyly of the Protozygoptera and their exact relationships with Archizygoptera remain uncertain. The present study of new fossil wings of Protozygoptera from the Upper Permian of France adds some information for the phylogenetic study of this suborder. [Palaeontology, Vol. 42, Part 1, 1999, pp , 1 pl.] The Palaeontological Association

2 84 PALAEONTOLOGY, VOLUME 42 TEXT-FIG. 1. Location of the insect sites F23 (¼ Les Canals) and F31 (¼ Bouisset) in the southern and western parts of the Lodève Permian basin. ENTOMOLOGICAL SITES 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; Laversanne 1976; Alabouvette et al. 1982; Odin 1986; Gand 1987; Châteauneuf and Gand 1989). Sites F23 and F31 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 Permian comprises two sedimentary cycles separated by the Rabejac conglomerate. Within the second cycle, the TEXT-FIG. 2.A, the Lodève Permian, after Odin (1986). Fo ¼ formations; Ens ¼ Members: LI ¼ Lower Loiras, LS ¼ Upper Loiras, MA ¼ Mas d Alary, VI ¼ Lower Viala, VS ¼ Upper Viala; Cogema profile with V.A. ¼ volcanic ashes in Roman numerals; En ¼ environments. B, section of the Saxonian with insect levels F23 Les Canals, with Epilestes gallica sp. nov., and F31 ¼ Bouisset, with Lodevia longialata gen. et sp. nov. Odin ¼ Odin 1986; R ¼ bed-marks; a ¼ insect wings, c ¼ conchostracans, g ¼ gastropods, i ¼ Isopodichnus sp., p ¼ invertebrate tracks, t ¼ burrows, ti ¼ triopsids, v ¼ footprints. C, F29 up to F20 ¼ new fossiliferous levels in the Saxonian.

3 NEL ET AL.; PERMIAN INSECTS 85

4 86 PALAEONTOLOGY, VOLUME 42 TEXT-FIG. 3. Fossiliferous site F19; perspective view of lens I due to the filling of a gutter; j ¼ bedding-plane and l ¼ fossiliferous beds. Salagou F5 Formation containing the insects is the thickest, attaining almost 2000 m in the south of the basin. It is very monotonous and formed from multiple negative binary sequences of decimetric to decametric claystone and siltstone strata. Within the claystone small decimetric levels of siltstones sometimes 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 the sedimentology, these fossiliferous beds have been interpreted by Gand et al. (in press a, in press b) as representing crescent-shaped megaripples in temporary water bodies belonging to a semi-arid playa or pediplain environment. The Arièges site (¼ F19), as an example, comprises two lenses of very hard, red siltstones, laminated with carbonate cement and separated vertically by some ten metres. Each lens terminates upwards with a level of desiccation cracks acting as markers in the formation. That of lens I can be traced for a hundred metres but that of lens II is lost in the claystones after some ten metres of lateral extension. The dimensions of I and II are respectively 4 m and 6 m length by 0 5 m and 0 3 m maximum thickness. Text-figure 3 shows a cross section through lens I. Several sequences of poorly sorted siltstones can 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-references considered. 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 conglomerate allowed the Permian to be subdivided into Autunian and Saxonian Thuringian (Feys and Greber 1972). 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 by Doubinger et 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 yielded Supaia sp. which does 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 Russian Permian stratigraphy, make it possible to suggest the age of these beds at Arièges, Canals and Bouisset as lowermost Kazanian.

5 NEL ET AL.; PERMIAN INSECTS 87 SYSTEMATIC PALAEONTOLOGY Suborder PROTOZYGOPTERA Tillyard, 1925 Type family. Kennedyidae Tillyard, Phylogenetic position. Pritykina (1981, p. 34) considered that Protozygoptera and Archizygoptera Handlirsch, 1906 (Promomyrmeleontidae) should be put together in the infraorder Kennedyomorpha Pritykina, Carpenter (1992) concurred but renamed this group Archizygoptera. Nel and Henrotay (1992) considered that Protozygoptera requires revision and is probably not directly related to the Protomyrmeleontidae. Thus, these authors restricted Archizygoptera to the sole family Protomyrmeleontidae. Bechly (1995) also considered that the relative phylogenetic positions of true Odonata, Protozygoptera and Archizygoptera remain unresolved. Bechly (1996), however, proposed a new phylogenetic classification of these taxa, including Archizygoptera in Protozygoptera as sister-group of Permagrionoidea, Protozygoptera being the sister-group of Panodonata, viz. more Recent Odonatoptera. Family PERMOLESTIDAE Martynov, 1932 Type genus. Permolestes Martynov, 1932 (type species Permolestes gracilis Martynov, 1932; Kazanian, Upper Permian, Arkhangelsk District, Siberia). Other genera. Epilestes Martynov, 1937 (type species Epilestes kargalensis Martynov, 1937; lowermost Kazanian, Permian, Kargala, Semenovski, Siberia), Scytolestes Martynov, 1937 (type species Scytolestes stigmalis Martynov, 1937; Permian, Kargala, Semenovski, Siberia) and Solikamptilon Zalessky, 1948 (type species Solikamptilon remuliforme Zalessky, 1948; Permian, Sibera). Carpenter (1992) included the genus Sushkinia Martynov, 1930 (type species S. parvula Martynov, 1930 and S. elongata Martynov, 1930 from the Ufimian Horizon, Kungurian, Permian, Tikhie Gory, Kama River, Kazan, Russia) which was originally considered by Martynov (1930) as Kennedyidae. The attribution of Sushkinia to Permolestidae is uncertain because only the distal parts of the wings of both S. parvula and S. elongata are known, and so the wing venational synapomorphies of Permagrionoidea (¼ Permolestidae þ Permagrionidae) are unknown in these taxa. Furthermore, S. parvula and S. elongata share with Kennedyidae two synapomorphies, viz. no more than one crossvein beneath the pterostigma, and few postnodals. Thus, our present state of knowledge strongly suggests that Sushkinia is more closely related to Kennedyidae than to Permolestidae. Genus EPILESTES Martynov, 1937 Type species. Epilestes kargalensis Martynov, Epilestes gallica sp. 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, Musée Fleury, Lodève. Type locality and horizon. Upper Permian, Kazanian, Les Canals (¼ F23), Lodèvois, France. Diagnosis. The fore- (?) wing of this species is characterized as follows: long but rather wide, well petiolated; cubital area broad, with three rows of cells between CuA and posterior wing margin; CuA with numerous long branches; main veins are long and straight but without intercalary longitudinal veins

6 88 PALAEONTOLOGY, VOLUME 42 TEXT-FIG. 4. Epilestes gallica sp. nov.; holotype, Ld LAP 110, Fleury Museum in Lodève; wing venation. Scale bars represent 2 mm. between them; MP separated from Cu at wing base; pterostigma long and narrow; wing apex acute; nine postnodal 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 viceversa. No trace of coloration preserved. Distance from base to nodus 11 2 mm, from nodus to apex 17 7 mm. Nodus in a basal position. Distance from base to arculus 4 6 mm. Petiolus long and narrow, 5 3 mm long and 1 3 mm wide. MP þ Cu and AA þ AP strongly approximate in petiolus. CuP not preserved. MP basally separated from Cu, closely parallel 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 mm long and strongly convex, looking like a posterior branch MAb of MA (Nel et al. 1993) distally closing an anteriorly open 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, RP 00, IR2, RP2, and IR1 simple and nearly straight. CuA reaching posterior margin at 77 per cent., MP at 85 per cent. and MA at 89 per cent. of wing length. RP 00 beginning 7 1 mm distal of arculus and 0 5 mm distal of subnodus, IR2 1 9 mm distal of base of RP 00, 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, RP 00, IR2, RP2, IR1 and RP1. Wing apex acute. No oblique crossvein O between IR2 and RP2. Although nodus poorly preserved, nodal crossvein Cr between ScP and RA very clear and subnodus Sn distinctly oblique. Pterostigma poorly preserved but long and narrow, c. 3 mm long. No oblique pterostigmal brace. Postnodal area poorly preserved, but with numerous postnodal crossveins, not aligned with postsubnodal crossveins between RA and RP1. Antenodal area poorly preserved, 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, basal of subnodus. AA completely fused with AP. Opposite MAb, A strongly concave and very short oblique crossvein m-cu (sensu Bechly 1996), 0 1 mm long, between MP and CuA, and looking like a posterior branch of MP rather than other convex 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 the Protozygoptera: Permolestidae Epilestes, Scytolestes and Permolestes. The differences between these genera and Epilestes gallica sp. nov. are as follows. There is no secondary longitudinal vein between all

7 NEL ET AL.; PERMIAN INSECTS 89 main veins, unlike Permolestes gracilis which has one between RP2 and IR2 and between RP 00 and MA. MA of P. gracilis is distally zigzagged and indistinct in the area between RP 00 and MP. IR1 is straight and RP2 is not secondarly branched on IR2, unlike Scytolestes stigmalis. The wing of E. gallica sp. nov. is somewhat smaller (29 mm long) than that of E. kargalensis (31 mm long); there are three rows of cells in the cubital area instead of two in E. kargalensis; the wing apex is more acute than in E. kargalensis; and there are fewer subpostnodal crossveins. All other venational characters are identical in the two species. Martynov (1932) noted that the hindwing pterostigma of P. gracilis is distinctly shorter and in a more distal position than that of the forewing. From this remark, it is possible to infer that the holotype specimen of E. gallica might be a forewing. Phylogenetic affinities within Protozygoptera. Bechly (1996) considered Protozygoptera to be the sister group of Panodonata (¼ Tarsophlebioidea þ [Zygoptera þ Anisozygopteria]): the sister group of (Protozygoptera þ Panodonata ¼ Stigmoptera) being the Triadophlebiomorpha. Bechly s two main synapomorphies of Stigmoptera, absent in Triadophlebiomorpha, are: (1) the distal free part of vein CuP is secondarily absent (suppressed or fused with the anal vein) and (2) a sclerotized pterostigma is present. These synapomorphies are present in E. gallica. Bechly (1996) proposed the following synapomorphies for the Protozygoptera (¼ [Permagrionidae þpermolestidae] þ Archizygoptera): (1) strongly petiolated wings (convergent to fossil Triadophlebiomorpha and many Zygoptera) with a very long petiolus ; (2) nodus shifted in a more basal position (than in Triadophlebiomorpha) ; (3) regressive development 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 (including E. gallica) share characters (1) to (4) with other Protozygoptera but character (5) is figured only in P. gracilis. Martynov (1937) and Zalessky (1948) did not mention this vein in E. kargalensis, S. stigmalis and S. remuliforme. Character (5) is also unknown in E. gallica sp. nov. Furthermore, the new species shares with Protozygoptera the separation between CuA and MP distinctly basal of MAb (plesiomorphy). Within Protozygoptera, Permagrionoidea (¼ Permagrionidae þ Permolestidae) are characterized by the synapomorphy, shared by E. gallica, of nearly aligned arculus and AX2 (convergently acquired by some Zygoptera). Archizygoptera (sister group of Permagrionoidea within Protozygoptera) is characterized by the nodal and subnodal crossveins being transverse, non-aligned and reduced. This reduction of the nodal structures is absent in E. gallica. Following Bechly (1996), the Permagrionidae (including the Upper Permian type genus and species Permagrion falklandicus Tillyard, 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 M suppressed ; (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 following discussion, we list their possible synapomorphies. 1. The polarity of character (2) (pterostigmal structures) is uncertain, because the pterostigma is absent in Triadophlebiomorpha, sister group of Stigmoptera. The very long pterostigma of Permolestidae, with its non-oblique basal margin and its distal margin distinctly oblique, is a very peculiar structure, also present in Kennedyidae and Bakteniidae (Archizygoptera; Bechly 1996). The pterostigmal structure of Permagrionidae (basal and distal margins oblique and parallel) is shared by Archizygoptera: Protomyrmeleontidae and Panadonata. If it is supposed that the pterostigmal structure of Permagrionidae is an apomorphy, it would imply that it has been acquired convergently by Protomyrmeleontoidea and Panodonata. On the other hand, if the pterostigmal structure of Permolestidae is an apomorphy, it would also imply two convergences with Kennedyidae and Bakteniidae. Thus, it is still impossible to decide which of the two solutions is the more parsimonious. 2. The base of CuA is nearly opposite the arculus in Permagrion Tillyard, 1928, unlike in Permolestidae and Epilestes gallica sp. nov. However, as a situation similar to that in Permolestidae occurs in Triadophlebiidae (Triadophlebiomorpha) and Kennedyidae, it is probably a symplesiomorphy of Permolestidae. 3. Veinlet m-cu is aligned with MAb in Permagrion and E. gallica and distal of MAb in Permolestes. This structure was not mentioned by Martynov (1937) and Zalessky (1948) for other Permolestidae

8 90 PALAEONTOLOGY, VOLUME 42 (E. kargalensis, S. stigmalis and S. remuliforme). Thus, it is very difficult to determine the exact value of this character. Furthermore, the triadophlebioidean Triadophlebia madygenica Pritykina, 1981 shares with Permagrion the alignment of m-cu with MAb, suggesting that it is a plesiomorphy. 4. All Permolestidae, except perhaps E. kargalensis, have distinctly fewer postsubnodal than postnodal crossveins (three against five for P. gracilis; three against six for S. stigmalis; three against nine for E. gallica). The same situation occurs in Kennedyidae. The postnodal and subpostnodal areas of E. kargalensis seem to be poorly preserved, as several postnodal crossveins that should be present are not figured and several postsubnodals are indicated as dotted lines by Martynov (1937). There are as many postsubnodal crossveins as postnodal crossveins in Permagrion. The same situation occurs in Triadophlebioidea and Protomyrmeleontoidea (¼ Bakteniidae and Protomyrmeleontidae) and Panodonata. Thus, it is more likely that the reduction of the number of postsubnodal crossveins is a synapomorphy of Permolestidae, which is also present in Kennedyidae (synapomorphy or convergence?). As E. gallica has none of the strong apomorphies of Permagrion (characters (1) and (3) listed above) and as it shares one possible synapomorphy of the Permolestidae, we propose to include it in this family. It will be possible to verify this hypothesis after revision of the holotype of Epilestes kargalensis. Family PERMEPALLAGIDAE Martynov, 1938 Type genus. Permepallage Martynov, 1938 (type species Permepallage angustissima Martynov, 1938, from the Permian, Kazanian, Sojana River, Arkhangelsk district, Russia). Other species. Rohdendorf et al. (1961, p. 89, text-fig. 37) described a Permepallagidea fam., gen., sp. from the Permian of Russia, which is an incertae sedis because it is based on a poorly preserved costal portion of a wing. Wing venational autapomorphies (modified from Bechly 1996). Wings extremely slender; secondary antenodals present (possible reversal?); very long and concave oblique veinlet m-cu; postnodal and postsubnodal crossveins very numerous. Genus LODEVIA gen. nov. Derivation of name. From the town Lodève (Hérault, France). Types species. Lodevia longialata sp. nov. Diagnosis. The wing venation of Lodevia gen. nov. is similar to that of Permepallage, differing in: only one concave basal crossvein between MP and CuA, instead of two; absence of intercalary veins between branches of RP; obliquity of the subnodus, instead of being nearly perpendicular to RA as in Permepallage; obliquity of MAb directed towards wing apex, instead of being directed towards base as in Permepallage. The presence of numerous antenodal crossveins, considered by Bechly (1996) as an apomorphy of Permepallage, is unknown in Lodevia. Lodevia longialata sp. 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, Musée Fleury, Lodève. Type locality and horizon. Upper Permian, Kazanian, holotype from Bouisset (¼ F31), paratype from Le Moural D (¼ F21D), Lodèvois, France. Diagnosis. As for genus (this is the only recognized species).

9 NEL ET AL.; PERMIAN INSECTS 91 TEXT-FIG. 5. Lodevia longialata gen. et sp. nov.; holotype, Ld LAP 111, Fleury Museum in Lodève; wing venation. Scale bar represents 2 mm. Description Holotype (Text-fig. 5). Impression of a nearly complete wing very long and slender, about 38 mm long and 4 9 mm wide, which is only broken opposite the nodus. No trace of coloration preserved. Distance from base to nodus about 15 mm, from nodus to apex 23 mm. Nodus in a basal position. Distance from base to arculus 6 8 mm. Petiolus long and very narrow, 6 5 mm long and 1 0 mm wide. MP þ Cu and AA þ AP long parallel but not strongly approximate in petiolus. Transverse CuP not visible due to anal area covered by sediment, but CuP with no distal free part. MP not basally separated from Cu. MP þ Cu strongly concave. RP separated from MA by a long distance, 0 7 mm distal of base of the strongly oblique arculus. First crossvein MAb between MA and MP strongly convex and distally closing a basally open discoidal space. MA and MP strongly approximate and diverging again distally, thus seeming to close the discoidal 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, RP 00, IR2, RP2 and IR1 all nearly straight. CuA reaching posterior margin at 70 per cent., MP at 84 per cent. and MA at 90 per cent. of wing length. RP 00 probably beginning slightly distal of nodus. IR2 beginning about 4 mm distal of base of RP 00, RP2 6 2 mm distal of that of IR2. and IR1 beginning 4 0 mm distal of that of RP2. IR1 9 7 mm long. One row of cells in the areas between the main veins, i.e. posterior margin, CuA, MP, MA, RP 00, IR2, RP2, IR1 and RP1. Wing apex rounded. One oblique crossvein between IR2 and RP2, five cells distal of base of RP2. Nodus not preserved. Pterostigma long and narrow, 3 7 mm long, covering five cells, with its proximal side making a right angle with RA and its distal side oblique. No oblique pterostigmal brace. Postnodal area wellpreserved 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. No visible crossvein in area between RA and RP, basal of subnodus. AA completely fused with AP. A strongly concave and 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, RP 00, IR2, RP2 and IR1, all nearly straight with one row of cells in the areas between them. Nodus well preserved with the subnodus distinctly oblique and not aligned with nodal crossvein Cr. Postnodal area well-preserved with 16 visible postnodal crossveins, not aligned with the 15 postsubnodal crossveins between RA and RP1. Two visible crossveins in area RA and RP, basal of subnodus. RP 00 begins one cell distal of the subnodus. Remarks. We attribute these two specimens to the same species because they share the same organization of the main veins in the median part of the wing and the absence of any intercalary veins between them. Lodevia longialata gen. 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 strongly petiolated with AA and AP completely fused, petiolus very long, nodus shifted in a basal position. Its position within Protozygoptera is more difficult to establish. Nevertheless, it can be provisionally related to the Permepallagidae because of the unique synapomorphies within Protozygoptera: principally, numerous postnodal and subpostnodal crossveins, a very long and slender wing and oblique m-cu very long (distinctly longer than MAb). Lodevia gen. nov. and Permepallage also share the presence of numerous crossveins beneath the pterostigma between RA and RP1 (probable symplesiomorphy), the base of RP 00 one cell distal of the subnodus and the presence of numerous crossveins between RA and RP basal of the subnodus. Nevertheless, this attribution is still open because the family characters of Permepallagidae are not at all clear. Martynov (1938, p. 50) considered the presence of a basal enigmatic longitudinal concave (?) vein in the petiolus, fused with the anal margin about midway of the petiolus, and that of a small crossvein, between MP þ Cu and the previous enigmatic vein, as the main diagnostic character of Permepallage. As it is

10 92 PALAEONTOLOGY, VOLUME 42 TEXT-FIG. 6. Lodevia longialata gen. et sp. nov.; paratype, Ld LAP 424, Fleury Museum in Lodève; wing venation. Scale bar represents 2 mm. possible to homologize the small crossvein as CuP, because of its position, then the basal enigmatic longitudinal 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 was misinterpreted by Martynov, or it is only an artefact. Re-examination of the holotype of Permepallage angustissima will be necessary to solve this problem. Nevertheless, it is not possible to define Permepallagidae using this dubious character, which is, anyway, poorly preserved in the holotype of Lodevia longialata gen. et sp. nov. Besides, the presence of intercalary longitudinal veins between the branches of RP, which is considered by Bechly (1996) as an apomorphy of Permepallage, is not sufficient to exclude Lodevia from Permelpallagidae on account of similarities in Recent Zygoptera: Coenagrionidae. In them, the number of intercalary veins varies greatly through the different genera so that this character is of little significance at family level. Thus, including Lodevia in a new family is not necessary, and also because it would add no information to the relationships between it and Permepallage. CONCLUSIONS Previous to the description of Epilestes gallica sp. nov. and Lodevia longialata gen. et sp. nov., Arctotypus verneti Laurentiaux-Vieira and Laurentiaux, 1963 (Meganeuridae Tupinae, Upper Permian, Alpes Maritimes, France), was the only described odonatopteran from the Upper Permian of France. As undescribed Tupinae are also present in the Lodève s Formation, together with a new undescribed family, related to the Protanisoptera, the odonatan fauna from the Saxonian of France appears to have been abundant and diverse (Gand et al. in press b; Nel et al. in press). The two newly described taxa are attributed to the families Permepallagidae and Permolestidae which have been, until now, described only from the Upper Permian (Kazanian) of Russia. Thus, it is possible to infer 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 (sensu Huelsenbeck 1994) with the phylogeny proposed by Bechly (1996) (see Table 1 and Text-fig. 7). Their sister-group relationships 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 be supposed. The inferred Kazanian age needs further evidence based on the study of the other insects and their comparison with the entomofaunas of the Permian of Russia and North America. EXPLANATION OF PLATE 1 Fig. 1. Lodevia longialata gen. et sp. nov.; holotype, Ld LAP 111, Fleury Museum in Lodève; Site F31 ¼ Bouisset, Salagou Formation; 3 5. Fig. 2. Epilestes gallica sp. nov.; holotype, Ld LAP 110, Fleury Museum in Lodève; Site F23 ¼ les Canals, Salagou Formation; 4 25.

11 NEL et al., Lodevia, Epilestes PLATE 1

12 94 PALAEONTOLOGY, VOLUME 42 TABLE 1. Permo-Triassic taxa attributable to Protozygoptera. PERMAGRIONOIDEA PERMOLESTIDAE Martynov, 1932 Permolestes Martynov, 1932 Permolestes gracilis Martynov, 1932 (Upper Permian) Epilestes Martynov, 1937 Epilestes kargalensis Martynov, 1937 (Upper Permian) Epilestes gallica sp. nov. (Upper Permian) Scytolestes Martynov, 1937 Scytolestes stigmalis Martynov, 1937 (Upper Permian) Solikamptilon Zalessky, 1948 Solikamptilon remuliforme Zalessky, 1948 (Permian) PERMAGRIONIDAE Tillyard, 1928 Permagrion Tillyard, 1928 Permagrion falklandicus Tillyard, 1928 (Upper Permian) ARCHIZYGOPTERA KENNEDYIDAE Tillyard, 1925 Kennedya Tillyard, 1925 Kennedya mirabilis Tillyard, 1925 (Carpenter 1931, 1933, 1939, 1947) (Lower Permian) Kennedya tillyardi Carpenter, 1939 (Permian) Kennedya reducta Carpenter, 1939 (Permian) Kennedya fraseri Carpenter, 1947 (Permian) Kennedya carpenteri Pritykina, 1981 (Triassic) Kennedya gracilis Pritykina, 1981 (Triassic) Kennedya sp. (Tasch and Zimmerman 1959, 1962) (Permian) Progoneura Carpenter, 1931 Progoneura minuta Carpenter, 1931 (Permian) Progoneura nobilis Carpenter, 1947 (Permian) Progoneura venula Carpenter, 1947 (Permian) Progoneura sp. (Carpenter, 1947) (Permian) Suschkinia Martynov, 1930 Suschkinia parvula Martynov, 1930 (Permian) Suschkinia elongata Martynov, 1930 (Permian) Opter Sellards, 1909 Opter brongniarti Sellards, 1909 (Permian) PERMEPALLAGIDAE Martynov, 1938 Permepallage Martynov, 1938 Permepallage angustissima Martynov, 1938 (Upper Permian) Lodevia gen. nov. Lodevia longialata sp. nov. (Upper Permian) Permepallagidea fam., gen., sp. (Rohdendorf et al. 1961) (Upper Permian) PROTOMYRMELEONTOIDEA BATKENIIDAE Pritykina, 1981 Batkenia Pritykina, 1981 Batkenia pusilla Pritykina, 1981 (Triassic) Terskeja Pritykina, 1981 (Protomyrmeleontidae sensu Pritykina 1981; uncertain position after Nel and Henrotay 1992; in Batkeniidae after Bechly 1996) Terskeja paula Pritykina, 1981 (Triassic) Terskeja pumilio Pritykina, 1981 (Triassic) Terskeja tenuis Pritykina, 1981 (Triassic) Voltzialestes Nel, Papier, Grauvogel-Stamm and Gall, 1997 Voltzialestes triasicus Nel, Papier, Grauvogel-Stamm and Gall, 1997 (Lower Triassic) PROTOMYRMELEONTIDAE Handlirsch, 1906 Tillyardomyrmeleon Henrotay, Nel and Jarzembowski, 1997 Tillyardomyrmeleon petermilleri Henrotay, Nel and Jarzembowski, 1997 (Middle Triassic) Triassagrion Tillyard, 1922 Triassagrion australiense Tillyard, 1922 (Upper Triassic)

13 NEL ET AL.; PERMIAN INSECTS 95 TEXT-FIG. 7. Phylogenetic tree of the families of Protozygoptera (after Bechly 1996), with oldest records. Acknowledgements. This work is a contribution to the team Paléontologie et Biostratigraphie du Permien of the UMR/CNRS Paléontologie analytique et Géologie sédimentaire. It is supported by the UMR 5561/CNRS. Our great thanks to Mr Godon and Mme Bussière of the Centre des Sciences de la Terre de l Université de Bourgogne, for the photographs of the fossils and the preparation of the figures. REFERENCES ALABOUVETTE, B., AUBAGNE, M., BAMBIER, A., FEIST, R. and PALOC, H Notice explicative de la feuille Lodève à 1/ Bureau de Recherches Géologiques et Minières, Orléan-La-Source, 52 pp. BECHLY, G Morphologische Untersuchungen am Flügelgeäder der rezenten Libellen und deren Stammgruppenvertreter (Insecta, Odonatoptera), unter besonderer Berücksichtigung der Phylogenetischen Systematik und des Grundplanes der Odonata. Petalura, Special Volume, 1, Morphologische Untersuchungen am Flügeäder der rezenten Libellen und deren Stammgruppenvertreter (Insecta, Pterygota, Odonata), unter besonderer Berücksichtigung der Phylogenetischen Systematik und der Grundplanes der Odonata. Petalura, Special Volume, 2, CARPENTER, F. M The Lower Permian insects of Kansas. Part 2: the orders Palaeodictyoptera, Protodonata, and Odonata. American Journal of Science, Series 5, 21, The Lower Permian insects of Kansas. Part 6, Delopteridae, Protelytroptera, Plectoptera and a new collection of Protodonata, Odonata, Megasecoptera, Homoptera and Psocoptera. Proceedings of the American Academy of Arts and Sciences, 68, The Lower Permian insects of Kansas. Part 8, additional Megasecoptera, Protodonata, Odonata, Homoptera Psocoptera, Plecoptera, and Protoperlaria. Proceedings of the American Academy of Arts and Sciences, 73,

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