Corresponding author: Jakub Prokop (
Academic editor: D. Shcherbakov
Three new palaeopteran insects are described from the Middle Permian (Guadalupian) of Salagou Formation in the Lodève Basin (South of France), viz. the diaphanopterodean
Before the tremendous effort of collect of Dr Jean Lapeyrie who brought together a large collection of thousands of fossils, the ‘red’ Middle Permian Salagou Formation (Lodève Basin) was considered devoid of fossils. Palaeoenvironment of Lodève Basin could be characterized as climate controlled playa with ephemeral pools colonized by aquatic clam shrimps (Conchostraca) and tadpole shrimps (Notostraca) together with insects (
We describe herein four new fossil insects from the Salagou Formation those correspond to important new taxa. Unfortunately the great majority of this entomofauna is represented by isolated wings. The insect bodies have been destroyed by the abundant necrophagous animals (mainly
The material described in the present paper comes from the Middle Permian deposits of Salagou Formation found in several localities in the Lodève Basin, Hérault, France (
The material was observed under stereomicroscope Olympus SZX-9 and venation pattern drawings were drawn directly through stereomicroscope by camera lucida. Photographs were made using digital camera Nikon D80 with macro lens Nikon AF-S VR Micro-Nikkor 105 mm by single sided cross-light exposure.
We follow the wing venation nomenclature of
Specimen Ld LAP 483 (Lapeyrie collection, imprint of proximal part of forewing), stored at the Musée of Lodève, France.
Middle Permian, Guadalupian, Lodève Basin, Salagou Formation, Lodève, Hérault, France (
Counter-imprint of a fore wing without trace of preserved coloration, strongly developed corrugation of longitudinal veins; basal part of fore wing 22.7 mm long and 11.6 mm wide, estimated total length about 45 mm; area between ScP and C rather broad with numerous simple cross-veins; concave ScP straight and basally running close to radial and medial veins; RA nearly straight; RA and RP extremely approximate between wing base and a point situated 17.5 mm distally at which RP strongly diverges from RA; convex MA diverging from MP nearly at right angle and directed towards radial veins, 11.3 mm from wing base; MA distally closely parallel to RA and RP for 5.4 mm, then nearly touching RP at one point; RP emerging 6.4 mm distal of base of MA; concave MP nearly straight; convex CuA diverging from CuP close to wing base and running parallel to medial veins, CuA with two visible terminal branches; simple CuP strongly concave and only weakly curved; anal area partly preserved, first anal vein of neutral polarity ending with two main branches, second anal vein convex and distally pectinate with several branches connected by rather dense network of cells; along posterior wing margin a broad area between CuP and first anal vein; a small pentagonal elongate cell below second anal vein, near wing base.
Genus and species indet.(
This fossilbears a combination of the main characters of ephemeropterid family
Wing venation only. ScP ending on anterior wing margin near mid part of wing; stems of CuA, M and R very close; CuA, MP and MA diverging at the same point; MA and radial stem very closely parallel; MA distally fused for a short distance with RP; a very broad area between RA and anterior wing margin with several long simple oblique crossveins; a very broad area between CuA and CuP; CuA with weak secondary posterior branches.
Named after Prof. Alexandr Rasnitsyn.
That of the family.
Holotype LdLAP 318A (Lapeyrie collection, prints of two identical wings), stored at the Musée of Lodève, France.
Middle Permian, Guadalupian, Mérifons Member, Salagou Formation, Lodève, Languedoc, France (
That of the family.
Wing 11.8 mm long, 3.9 mm wide; ScP simple, ending in costal margin near mid part of wing, a narrow area between it and C without visible crossveins; area between C and RA relatively broad, 0.6 mm wide, with a row of simple oblique crossveins; RA simple ending on anterior wing margin 0.4 mm from wing apex; RP separating from RA 2.6 mm from wing base; RP with six-seven branches and covering a broad area with series of crossveins; CuA, MP, and MA diverging at the same point, 1.3 mm from wing base; MA very closely parallel to radial stem and reaching RP, fused with it for 0.6 mm, and separating again distally, MA simple, slightly curved; MP simple; CuA with three short distal branches; a series of crossveins between MP and CuA; a broad area between CuA and CuP with crossveins; one anal vein preserved at least.
As the two wings show the same convexity of the veins and are very close, they are likely to be a fore- and a hindwing of the same specimen, but it is impossible to determine which one is the forewing.
Note that the Upper Carboniferous monotypic family
The other structures of wing venation are similar in
Named after the Permian and the genus
Wing venation only. ScP long, terminating to C close to wing apex; absence of ‘trifurcation between R, M, and CuA near wing base; CuA never touching M; MA and RP never touching.
Holotype Ld LAP 365ab (Lapeyrie collection, imprint and counterimprint of a complete wing), stored at the Musée of Lodève, France.
Middle Permian, Lodève Basin, Salagou Formation, Les Vignasses, Lodève, Hérault, France (
Complete wing without trace of coloration; wing 11.7 mm long and 3.3 mm wide; ScP simple and parallel to RA ending on costal margin close to wing apex (about 0.5 mm); stems R+M basally distinctly curved, with R and M separating about 1.8 mm from wing base; convex simple RA nearly straight, and ending in wing apex; RP separating from RA about 1/3 of wing length, RP with three long branches ending on posterior wing margin; simple convex MA shortly connected to RP by a crossvein; concave MP deeply forked 0.9 mm from division R and M; anterior branch of MP secondary bifurcated, posterior branch simple; simple convex CuA, basally close to R+M; no apparent ‘trichotomy’ R/M/CuA; simple concave CuP; anal area with three or four veins.
Named after Lodève town and Hymen.
Wing characters only. Presence of basal fusion of M with CuA; broad areas between C and ScP and between ScP and RA without apical ‘pterostigma’-like structure, RP with five posterior parallel branches; strong convexity of basally connected veins R, M, and CuA, forming a curve.
Holotype LdLAP 556 (imprint of nearly complete wing, basal part not well preserved, Lapeyrie collection), stored at the Musée of Lodève, France.
Middle Permian, Lodève Basin, Salagou Formation, Les Canals, Lodève Basin, Hérault, France (
Wing elongated and basally narrow with no trace of coloration preserved, wing fragment about 16.5 mm long, estimated length about 19.5 mm width 4.2 mm in widest part; anterior margin nearly straight, ScP straight ending to C close to wing apex; areas between costal margin and ScP and ScP and RA rather broad; stem of R basally connected to M diverging 1/3 of wing length; division of RA and RP about 8.5 mm from the wing base, RA simple and straight ending in wing apex; RP pectinate with five branches ending on posterior wing margin, crossveins organized in rows between branches of RP; simple MA strongly diverges from MP 1.2 mm from division of stems R and M, and well connected with RP for distance; MP, CuA separates from CuP near wing base, CuA and M basally fused and closely parallel to R, separating 3.0 mm from base of CuA; CuA and CuP simple and straight with a few crossveins between them; anal area reduced, one anal vein present close the wing base.
Nevertheless
The genus
We described in the present study two new representatives of
Authors are grateful to reviewers Nina Sinitshenkova (Palaeontological Institute RAS, Moscow), Nikita J. Kluge (Saint-Petersburg State University, St.-Petersburg) and journal editor Dmitry Shcherbakov (Palaeontological Institute RAS, Moscow) for their insightful comments and suggestions. First author thank to Dr. Susan Butts for access to the collection of Peabody Museum of Natural History Yale University (New Haven, USA) and possibility to take photograph of type specimen. We sincerely thank Dr Jean Lapeyrie and Stéphane Foucher from the Musée de Lodève for having made available for study the impressive collection of fossil insects from Lodève. The first author (JP) acknowledges the research support from the Grant Agency of the Czech Republic No. P210/10/0633 and Ministry of Education of the Czech Republic MSM 0021620828.