Archepseudophylla fossilis sp. nov. Other included species: Archepseudophylla laurenti (Théobald, 1937) comb. nov., and Archepseudophylla melanospila (Cockerel, 1921) comb. nov.

Named after Arche (in Latin) and Pseudophylla in reference to its leaf-like tegmina and antiquity.

Tegmina covered by a net of large polygonal cells, subdivided into smaller cells; subcostal area broad, but covering only 36% of wing width; ScP and radius strongly diverging at about 40% of wing length; RA, RP and median vein strongly zigzagged; RP and median vein distally vanishing in the net of large cells; apex of tegmina rounded.

Archepseudophylla fossilis sp. n. (Fig. 1, Fig. 2, Fig. 3, Fig. 4 and Fig. 5)

Holotype (Fig. 1, Fig. 2 and Fig. 3): tegminae elongate, with oval apex, 35.0 mm long, 13.8 mm wide; area between ScP and Costa very broad, 4.5 mm wide, but covering only 36% of whole wing width in its broadest part, divided into two rows of large quadrangular or pentagonal cells by secondary branches of ScP, these branches being zigzagged and these large cells subdivided into a net of smaller cells by veinlets; ScP and radius R closely parallel in basal 40% of wing length, but strongly diverging distally, with a row of five large pentagonal to heptagonal cells between them; ScP strongly zigzagged in its distal part, ending on costa 26.0 mm from the wing base and 10.7 mm from the wing apex; basal part of the radius straight; RP separating from RA 16.9 mm from the wing base; RA distally zigzagged, distally parallel to costa and ending on the posterior wing margin, 1.5 mm close to wing apex, area between RA and costa 1.7 mm wide near the wing apex; RP strongly zigzagged and rather indistinct in its distal part; area between RA and RP with two rows of very large quadrangular to octagonal cells, all of them being secondarily divided into smaller irregular cells by veinlets, but frequently with a central cell or group of very small cells; median vein straight at its very base and strongly zigzagged distally, more or less vanishing in a net of very large cells identical to those of the area between RA and RP; 3–4 rows of large cells in the area between the radius/RP and the posterior wing margin; a large cell at the wing base below the median vein; cubito-anal area not visible, probably fossilized inside the matrix, this structure being currently perpendicular to the main part of the wing in the Tettigoniidae.

The paratype specimens from Céreste (Fig. 4 and Fig. 5) have nearly the same tegmina venation. Their tegmina are covered by a net of dark spots located in the central part of the polygonal cells.

Holotype specimen MNHN-LP-R.63854 (Camoins-les-Bains), imprint and counterprint of a nearly complete tegmina, with only antero-basal angle missing; no trace of coloration preserved, paratypes from Céreste: specimens MNHN-LP-R.63885, R.63886, R.63887, other material: 11 other specimens, Laboratory of Palaeontology, National Museum of Natural History, Paris, France.

After the fossil state of this species.

Early Oligocene [9] and [12], Camoins-les-Bains, Marseilles Basin, Bouches-du-Rhône, France.

Early Oligocene, Céreste, Alpes-de-Haute-Provence, France.

The type specimen of L. laurenti from Célas (Gard, France) is lost, but it is clearly visible on the photograph in Théobald [16 (pl. 1, Fig. 1)] that it had a colour pattern nearly identical to that of the specimens from the Oligocene of Céreste. The reconstruction of the tegmina venation of Théobald [16 (p. 114, text-fig. 8)] is certainly erroneous, as at least its distal branches of RA and RP were zigzagged as in the material from Céreste (visible on the photograph). The tegmina veins of Lithymnetes guttatus are not zigzagged [13 (pl. 17, fig. 14)]. Therefore, L. laurenti has to be removed from the genus Lithymnetes and transferred to the genus Archepseudophylla gen. nov. as Archepseudophylla laurenti comb. nov. It belongs to the Tettigoniidae, and not to the Hagloidea: ‘Prophalangopseinae’ (contra [14]). A. laurenti differs from the material from Céreste by its smaller dimensions (tegmina length 25 mm, instead of 32 to 35.5 mm in the material from Céreste). Nel [6] already noted this difference, but considered that it was not sufficient for a species separation. At that time, he had only three specimens available from Céreste, but the same range of wing lengths is now observable on 15 specimens. This strongly supports a specific separation between the fossils from Célas and Céreste. There is no clear difference between the tegmina venation and dimensions of the specimens from Camoins and Céreste. The unique difference is the absence of spot pattern on the tegmina of the specimen from Camoins, but this is probably related to its preservation as an imprint without any trace of coloration preserved, and not a mummy of the animal itself as for the Céreste's material. Therefore we consider that they belong to the same species.

‘Poekilloptera’ melanospila Cockerell, 1921 from the Bembridge Marls, Isle of Wight, of Early Oligocene age, was originally placed in Hemiptera: Fulgoridae; however, Szwedo et al. [15] considered that it was not a homopteran (the recent genus Poekilloptera Latreille 1796 belongs to the Hemiptera: Flatidae). Szwedo (pers. comm.) suggested that it might be a plant fossil; however, Margaret Collinson (pers. comm.) examined it and confirmed it is not a plant. Specimen In.24356 at the Natural History Museum, London, is the holotype (Fig. 6). It consists of a fragment of wing, clearly showing the subdivided large polygonal cells with a central colour spot, characteristic of Archepseudophylla. Unfortunately, it is too fragmentary to work out the full length of the tegmen, so it is uncertain whether it is conspecific with A. fossilis or A. laurenti or constitutes a separate species. For the moment, it should be regarded as a separate species, Archepseudophylla melanopsila (Cockerell, 1921) comb. nov. One of us (A.R.) has found an additional specimen in the collections of the Natural History Museum, London, In.25803/In.17326 (the counterpart is only a fragment) (Fig. 7). The specimen consists of wings, thoracic segments, pronotum and antennal bases. The wings are crumpled and fragmentary, with the characteristic subdivided polygonal cell and central colour spot, though unfortunately the length of the tegmina cannot be ascertained. The polygonal cells are similar in size to those of the holotype of Archepseudophylla melanopsila, so this specimen should be regarded as conspecific. The thoracic segments and pronotum have a distinctive punctuate ornament, the pronotum is 3.7 mm wide and the basal antennal segment is 0.7 mm wide. It is hoped that more complete material of A. melanospila will be found in due course.

The tegmina of Archepseudophylla are clearly leaf-like, with a broad subcostal area, veins ScP and radius closely parallel and displaced close to the middle part of the wing, and a net of large polygonal cells, subdivided into smaller cells, looking like the cells of a large leaf. Such wings can be found in some recent Tettigoniidae of the subfamilies Phaneropterinae Burmeister, 1838 and Pseudophyllinae Burmeister, 1840 [10]. The current classification of the tettigoniid subfamilies is not based on tegmina venation, but on other body characters, not available in Archepseudophylla fossilis [1]. Nevertheless, affinities with Phaneropterinae are unlikely because of the strong divergence between ScP and RA in the distal part of wing, not shared by leaf-like Phaneropterinae (see [10]). Within the Pseudophyllinae, A. fossilis looks like some recent Neotropical Pterochrozini Brunner von Wattenwyl, 1895 (Pterochroza Serville, 1831), Pterophyllini (Scopioricus), and Australian and Madagascarian Simoderini Brunner von Wattenwyl, 1895 (Lonchitophyllum Brunner von Wattenwyl, 1895 or Phyrama Karsch, 1889). But we could not find a recent taxon with such a strongly zigzagged RP and with a relatively reduced subcostal area, covering only 36% of wing width.

The specimens of Archepseudophylla constitute the first fossil records of the tettigoniid subfamily Pseudophyllinae. Because of the presence of strong convergence of mimicry in the tegmina among the tettigoniid tribes and genera, it is not possible to ascertain its affinities within the Pseudophyllinae.