These fossils have been found by surface prospecting at several hundred meters of distance, near a site already known for footprints and invertebrates traces [15] and [46]. It is the fossiliferous site F7 of the Coulet-Redon hill, located to the north of Roquebrune-sur-Argens. The new sites belong to the upper part of the Pradineaux Formation (Fig. 1) overlying in this area by that of Mitan [46].

The F7n (F7 for the Collet-Redon fossiliferous site and n for Notostraca) having supplied triopsid carapaces is situated in the east side of the hill, close to it summit but as the plane bedded mudstone fossiliferous slabs were collected in a small ravine, we do not know exactly the corresponding layer. Nevertheless, clearly, this bed is located above the δ 2 dolerit, equivalent of the B3 trachy-andesit, this latter being described from the Esterel basin by Boucarut [13]. In this site F7n, some sheeted silstones show also large plants stem prints (Fig. 2).

The F7i (i for insect) site is settled on a slope of the “Petit Coulet”. Herein, large red laminites show ripple marks (Fig. 3) and plant imprints lightly displaced from the original beds by gravity. The wing fragment was found in this place by M. Butchlii above layers bearing the footprints Microsauripus acutipes Moodie, 1929 similar to those of Varanopus curvidactylus Moodie, 1929 but smaller (Gand in [16], [17] and [23]) (Fig. 4) and Dromopus didactylus Moodie, 1930 (Fig. 5). These ichnites were collected in the quarry by the third author [15]. Stromatolitic-like structures were also remarked by RG and LB (Fig. 6). All these fossiliferous beds underlay the local black δ 2 dolerite.

The Pradinaux Formation [33], [45], [46] and [47] is on average 100 m thick. It begins by conglomerates with many rhyolitic pebbles coming from the underlying A7 flow, dated from radio-isotopes of 272, 5 ± 0, 3 Ma [34] and [49]; for example, Upper Kungurian. The above following beds are fluvial greenish to beige-brownish sandstones. Overlying are finer and often sheeted sandstone/siltstones in which sedimentological, biological marks (ripples, rain-drops, dessication, footprints, burrows, invertebrate trackways, etc.) and macroflora are rather frequent [15] and [45] (Fig. 2). All meaning deposits in shallow water environment. In the Upper part of the Formation are green to brown lacustrine carbonate-rich mudstones, decimetric thick [45] and [47]. Some carbonate rolls are also met. They were made in marsh environment.

In the neighbor Esterel basin (Fig. 1), some of the carbonates beds, located above the B3 trachy-andesit flow, equivalent of the δ 2 dolerite of the Bas-Argens basin) supplied ostracods. From Lethiers et al. [32], these fossils allow to date Early Tatarian (= Midian) the fossiliferous beds; for example, Capitanian (265.8 to 260.4 Myr) sensu the global scale (ICS 2004) and sensu Menning et al. [34] (265 to 262 Myr). The Notostraca beds of the Pradineaux Formation would be Late Capitanian (Guadalupian) in age.

The triopsids belong to the following class, order and family respectively (Fig. 7 and Fig. 8),

Class Branchiopoda Latreille, 1817

Order Notostraca Sars, 1867

Family Triopsidae Keilhack, 1909

Material. Specimen MNHN-LP-R 63918, stored at the Muséum national d’histoire naturelle, Paris, France. A piece of siltstones (around 20 μm grain), 4 × 5 cm, 1 cm thick, with triopsid carapaces prints. In one face of the slab, imprints of carapaces are easily identifiable (Fig. 7), but there are also more poorly preserved prints and counterimprints on its back, possibly indicative of a multilayer deposit of carapaces.

The Fig. 7 shows the silstone slab with external cephalothoracic carapaces (or dorsal shield) of a dozen of individuals, some of them recovered with sediments. The reconstitution of this animal is proposed in Fig. 8. One can see that the ovoid-shaped shield is characteristic, with a streamlined dorsal carapace with prints of the strong lateral maxillary glands [25], [42], [43] and [44].

These fossils are attributed to notostracan Branchiopodae (Crustacea) of the family Triopsidae [42], which is known for its exceptional morphological stability from Carboniferous to Holocene, suggesting also a great ecological stability. The three best visible individuals (specimens a, b, c in Fig. 7) are about 9 to 11 mm long. The others being of the same length, we suppose that all the individuals were from the same cohort (same age). The distribution of these imprints on the slab, with at least three shields in dorsal position, and one concave imprint in ventral position, together with the absence of abdomen and telson suggest that they correspond to postmortem disarticulated bodies, deposited in a water depression.

We cannot assign these fossils to a precise triopsid species or genus because of the absence of diagnostic characters, all located on the telson or on the thoracic appendages. Gand et al. [25] and [26] described Lepidurus occitaniacus and Triops cancriformis permiensis from the Permian Lodève basin thank to the presence of thoracic-abdominal segments and supra-anal plates fossilised with carapaces.

We follow the wing venation nomenclature of Kukalová-Peck [30] and Willmann [48], with the reserves concerning the possible fusion and relative positions of CuP and AA1 + 2, already indicated in Prokop et al. [41].

Class Insecta L., 1758

Order Syntonopterida Handlirsch, 1911 and/or Superfamily Syntonopteroidea Kukalová-Peck, 1985

Family Syntonopteridae Handlirsch, 1911

List of included genera, after Prokop et al. [41]. Syntonoptera Handlirsch, 1911 (type species: S. schucherti Handlirsch, 1911 by monotypy); Lithoneura Carpenter, 1938 (type species: L. lameeri Carpenter, 1938); a new genus and species described by Prokop et al. [41]; Gallolithoneura gen. nov. (type species: Gallolithoneura butchlii sp. nov.)

Genus Gallolithoneura gen. n.

Type species. Gallolithoneura butchlii sp. n.

Derivation of name. Named after the Latin name Gallia for France, and Lithoneura.

Diagnosis. Apex of ScP probably near wing apex; forks of RP and MA at the same level, well distal of that of MP; fork of MP distal of that CuA; MP and CuA with single forks into two branches, with areas between them triangular in shape and rather reduced.

Gallolithoneura butchlii sp. n. (Fig. 9 and Fig. 10)

Description. Print of the median part of a wing with apex and base missing, no trace of coloration preserved; broadest part at about midwing; precostal area of CA and CP not preserved; simple concave ScP nearly parallel to anterior wing margin and reaching it distal forks of RP into branches, probably near wing apex; numerous straight and simple crossveins in area between ScP and C, not aligned with crossveins in area between ScP and RA; RA and RP probably fused a little basal of preserved part of wing; RA strong, convex, simple, parallel to ScP and ending probably in wing apex; R and MA strongly approximate in their basal parts; MP clearly concave, distally forked into two strong posterior branches, 8.4 mm basal of fork of MA, both branches of MP slightly curved posteriorly and ending on posterior wing margin well distal of level of fork of RP, area between branches of MP rather broad, with probably one secondary longitudinal vein; MA convex with two long branches and an intercalary veins between them; base of first posterior branch of RP opposite fork of MA, first branch of RP concave, apparently simple and weakly curved, at least in its preserved part; a convex and simple intercalary vein between branches of RP; CuA clearly convex, with a strong distal fork 4.1 mm basal of level of fork of MP, anterodistal branch of CuA parallel to posterior branch of MP, posterobasal branch of CuA the shortest and more curved than the other one; area between branches of CuA triangular with a secondary concave longitudinal vein in it; concave CuP simple, weakly curved; AA convex but only its apex is preserved; wing membrane between main veins with straight, simple crossveins perpendicular to main veins.

Dimensions. Length of wing fragment about 40 mm, probable total length about 65 to 70 mm; wing width about 15.4 mm. Wingspan probably around 15 to 16 cm.

Material. Holotype specimen MNHN-LP-R 63893 (median part of a single wing; supposed folded on another wing), on a small siltite slab, 1 cm thick (laboratoire de paléontologie, Muséum national d’histoire naturelle, Paris, France).

Derivation of name. Named after Mr Lorentz Butchli who found and donated the type specimen.

Age and outcrop. Permian, Capitanian, Capitanian (Upper Guadalupian), Pradineaux Formation, Petit Coulet Redon Hill, La Bouverie, Roquebrune-sur-Argens, Var, France.

Diagnosis. As for the family.

Discussion. The presence of only simple crossveins perpendicular to main longitudinal veins in the wing membrane (archaedictyon reduced and transformed into a regular polygonal meshwork of crossveins) excludes affinities with the palaeodictyopteroid orders and many Palaeozoic Ephemeroptera. The general pattern of wing venation would be compatible with an attribution to the Odonatoptera, but under this hypothesis, the vein that we call here “MA” would be in fact “IR2” and in that case, it would be distally forked, which is unlikely in all Palaeozoic Odonatoptera. Thus, an attribution to this order is to be rejected.

The preserved parts of the wing venation of Gallolithoneura are very similar to that of a Syntonopteridae Handlirsch, 1911. It especially shows a striking resemblance to Syntonoptera schucherti Handlirsch, 1911 (Upper Carboniferous of Mazon Creek, USA) in the relative positions of the forks of RP and MA nearly at the same level and those of MP and CuA in a distinctly more basal position [14]. The only differences concern the broader cubital area in Syntonoptera with four branches instead of two in our fossil. The new genus and species that Prokop et al. [41] recently described from the English Upper Carboniferous differs from our fossil in the less distant forks of MP and MA, MP with five branches, instead of two in our fossil, and CuA with four branches, instead of two [41]. The third syntonopterid genus Lithoneura Carpenter, 1938 (Upper Carboniferous of Mazon Creek, USA) differs from Gallolithoneura in the position of fork of MP distinctly basal of that of CuA, instead of being slightly distal in Gallolithoneura [48].