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Ann. soc. entomol. Fr. (n.s.), 2010, 46 (1–2) : 242-246ARTICLE

Lusitaneura covensis n. gen., n. sp., fi rst Caloneurodea from the Carboniferous of Portugal (Insecta: Pterygota: Panorthoptera)

Abstract. Lusitaneura covensis n. gen., n. sp., from the Late Carboniferous (Lower Stephanian C) of the Douro Basin (NW of Portugal), is the fi rst Portuguese representative of the Palaeozoic insect order Caloneurodea.

Résumé. Lusitaneura covensis n. gen., n. sp., premier Caloneurodea du Carbonifère du Portugal (Insecta : Pterygota : Panorthoptera). Lusitaneura covensis n. gen., n. sp. est le premier représentant portugais de l’ordre d’insectes paléozoïques des Caloneurodea. Il provient du Carbonifère supérieur (Stéphanien C inférieur) du Bassin du Douro (NW du Portugal).Keywords: nsecta, Caloneurodea, Late Carboniferous, Douro Basin, Portugal.

João P. Loureiro (1), Pedro Correia (2), André Nel (3)* & Ary Pinto de Jesus (4)

(1) Viana do Castelo, Portugal (2) Porto, Portugal

(3) CNRS UMR 7205, Muséum National d’Histoire Naturelle, CP 50, Entomologie, 45 rue Buff on, F-75005 Paris, France (4) Departamento & Centro de Geologia da Universidade do Porto, Portugal

* Corresponding author

E-mail: anel@mnhn.fr, johnyloureiro@live.com.pt, correia.p@live.com.pt, adelmar@fc.up.ptAccepté le 28 mai 2009

Carlos Teixeira developed the study of the Carboniferous insects from Portugal in the years

1939-1950. He described alone or in collaboration with the French palaeoentomologist Daniel Laurentiaux numerous fossils, mainly ‘blattodean’ taxa. In Teixeira (1946), is done the fi rst description of fossil insects from the Upper Stephanian outcrop of São Pedro da Cova (Douro Carboniferous Basin: São Pedro da Cova and Pejão Coalfi elds), i.e. a Palaeodictyoptera Homaloneura ribeiroi, a Blattinopsidae, and two ‘blattodean’ Phyloblattidae Phyloblatta fonsecai and P. carringtoni. Th e same author found numerous material in other Portuguese outcrops (brief summary in Laurentiaux & Teixeira 1948, 1958), but no further fossil insect was found at São Pedro da Cova. Th anks to recent fi eld researches, J. Loureiro and P. Correia discovered new fossil insects in São Pedro da Cova coalfi led outcrops. Th ese are a Palaeodictyoptera, several ‘Blattodeans’ and the fi rst record of the order Caloneurodea from Portugal, currently described in the present work. Th is order is ranging stratigraphically from the Westphalian (Upper Carboniferous) to the Kazanian (Upper Permian) of North America, Europe, and Siberia. Its discovery in the Portuguese Upper Stephanian is not

surprising as this order is already well known from the French classical locality of Commentry similar in age as Douro Basin.

Geological setting Th e Douro-Beira Carboniferous Trough (NW

of Portugal) occurs in the geological setting of the Central Iberian Zone of Iberian Massif, striking NW-SE, and extends approximately 85 km from Apúlia (North of Porto) to Mioma (NE of Viseu) (Lemos de Sousa & Wagner 1983a, Pinto de Jesus 2001, 2003). All along the Douro-Beira Carboniferous Trough there are several terrestrial Carboniferous outcrops with ages ranging from the Westphalian to the Lower Stephanian C. Although the age of the Carboniferous outcrops in the SE part of the Douro-Beira Carboniferous Trough are yet to be confi rmed in detail, a model of pull-apart basins in favourable relation with the important strike-slip sinistral component of the Douro-Beira Carboniferous Trough is strongly suggested (Pinto de Jesus 2001, 2003, Pinto de Jesus & Lemos de Sousa 2003). Douro Basin (Fig. 1) is the major Carboniferous basin present in the Douro-Beira Carboniferous Trough, forming a narrow strip, extended for 53 km from São Pedro Fins (East of Porto) until Janarde (East of Arouca). Sedimentary polarity indicates the base to be SW and the top towards NE.

Th e base of the Douro Basin rests unconformable over the Lower Cambrian (“Complexo Xisto-grauváquico”), and the top is cut by a high-angle reverse

A fi rst Caloneurodea from the Carniferous of Portugal

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fault that has placed the Early Palaeozoic formations of the reverse limb of the Valongo Anticline over the coal measures of the Douro Basin (Domingos et al. 1983; Lemos de Sousa 1984; Wagner et al. 1984; Pinto de Jesus 2001, 2003; Pinto de Jesus & Lemos de Sousa 2003).

Th e Douro Basin fi lling results from continental intramontane syntectonic sedimentation, with alluvial fans debris fl ow dominated at the base, and passing upwards to lacustrine/palustrine facies, truncated by a multistory-multichannel fl uvial braided system, and the top of the sequence showing lacustrine facies with deltaic features. Th e sequence is tectonically repeated in the tectonic slices resulting from the latest tectonic phase of variscan orogeny (Pinto de Jesus 1987, 2001, 2003; Oliveira & Pinto de Jesus 1998; Pinto de Jesus & Lemos de Sousa 2003).

Douro Basin is dated from the Lower Stephanian

C on the basis of palaeobotanical (Corsin & Lemos de Sousa 1972; Lemos de Sousa & Wagner 1983a, 1985; Wagner 1983; Wagner & Lemos de Sousa 1983, 1985), palaezoological (Eagar 1983), and palynological (Fernandes et al. 1997) studies.

For the megafl oristic studies, Lemos de Sousa & Wagner (1983b), put into special evidence the occur-rence of Pseudomariopteris cf. busqueti (Zeiller) Danzé-Corsin, of cf. Taeniopteris multinervis Weiss and of Neu-ropteris ovata var. pseudovata Gothan & Sze. Wagner & Lemos de Sousa (1983a) remarks on palaeogeography and conclusions for the intramontane sedimentation are based on the occurrence of extrabasinal elements comprising Ernestiodendron fi liciforme (Von Schlothe-im pars) Florin, cf. Lebachia frondosa var. zeilleri Florin and Lebachia parvifolia Florin, and also the presence of Dicranophyllum gallicum Grand’Eury and Dicrano-phyllum lusitanicum (Heer).

Figure 1Main geological regional setting of the Douro basin (lower Stephanian C; NW of Portugal), modifi ed from Pinto de Jesus (2001).

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From palaeozoological studies, Eagar (1983) concludes after the occurrence of Anthraconaia lusitanica (Teixeira) Eagar and of Anthraconaia (?) altissima Eagar.

Systematic palaeontologyWe follow the nomenclature of wing venation and

classifi cation of Béthoux et al. (2004).

Order Caloneurodea Handlirsch 1937

Family uncertain

Lusitaneura n. gen.

Type species. Lusitaneura covensis n. sp.

Etymology. Named after Lusitania and Caloneura.Diagnosis. Wing characters only. ScP reaching costal margin in second third of wing length and subcostal area not very broad; three posterior branches of RP; MA with two branches with anterior branch not making a strong curve, and posterior branch with small secondary twigs emerging from it; no vein distally emerging from MP+CuA+CuPaα; no small branches of CuPaβ near posterior wing margin.

Lusitaneura covensis n. sp.(Fig. 2)

Material. Holotype stored in private collection of João Loureiro, Viana do Castelo (a nearly complete wing, with cross-veins not well preserved except in anal area and between RA and RP). A plastic cast is stored at the Muséum national d’Histoire naturelle, Paris, France.Etymology. Named after São Pedro da Cova village.Age and outcrop. Upper Carboniferous, Lower Stephanian C, Douro Basin, São Pedro da Cova coalfi eld, Portugal.Description. A nearly complete wing, 21.7 mm long, 4.6 mm wide; ScP reaching anterior wing margin 15.1 mm distal of wing base; RA simple; fi rst posterior branch of RP 10.0 mm of its origin; RP posteriorly pectinate with three branches, all

simple; MA emerging from M+CuA+CuPaα just distal of the origin of RP; MA branched 1.2 mm distal of its origin; anterior branch of MA simple, posterior branch with three small posterior branches; MP+CuA+CuPaα simple, strongly convex; MP not emerging from MP+CuA+CuPaα; CuPaβ and CuPb simple and strongly concave; AA1 simple, weakly convex; area between AA1 and AA2 with intercalary veins; cross-veins not well preserved except in anal area and between RA and RP, simple and straight.

DiscussionLusitaneura n. gen. falls in the Caloneurodea

Handlirsch, 1937 because it has the diagnostic char-acters of this order: MA with two branches; a vein MP+CuA+CuPaα simple; fusion of CuPaα with MP+CuA; MP+CuA+CuPaα (convex) and CuPaβ (concave) close, parallel and straight; MP running fused with CuA+CuPaα at length; absence of anal fan. It has also the secondary character of the group, viz. RP posteriorly pectinated; marked convexity of cross-veins (visible only in some of them, between RA and RP; absence of anterior branch of RA. Th e very short anterior branch CuPaα between MP+CuA and CuPa, typical of the Panorthoptera is hardly visible because this part of the wing is deteriorated.

For the weakness of the current familial classifi cation within the Caloneurodea, we compare our fossil with the various genera currently attributed to this order (Béthoux et al. 2004).

Affi nities with Caloneura Brongniart 1885 is un-likely because the apex of ScP is near the second third of wing length and subcostal area is not very broad. Lusitaneura n. gen. diff ers from Gigagramma Béthoux et al. 2004 in the absence of any vein distally emerging from CuA+CuPaα, and in the more basal position of apex of ScP. Lusitaneura n. gen. diff ers from Sthena-rocera Brongniart 1885 in the presence of only three posterior branches of RP instead of 8–9. It diff ers from Apsidoneura Carpenter 1943 in the apex of ScP in a

Figure 2 Photograph of wing of Lusitaneura covensis n. gen., n. sp. (scale bar represents 10 mm)

A fi rst Caloneurodea from the Carniferous of Portugal

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more basal position, anterior branch of MA not mak-ing a strong curve (also diff erences with Homaloptila Handlirsch 1919), and RP with three branches, instead of two. Lusitaneura n. gen.diff ers from Pleisiogramma Carpenter, 1943 in the MA with two branches, instead of being simple, and the apex of ScP in a more basal position. Diff erences with Paleuthygramma Martynov 1930 are the same plus the presence of a rather broad area of RP, instead of being strongly reduced in the latter. Th e same last diff erence concerns Euthygramma Martynov, 1928 and Anomalogramma Carpenter 1943 and Nanogramma Béthoux et al. 2004. Th e Gelasop-teridae Carpenter 1976 (Gelasopteron Carpenter 1976) are Neoptera of uncertain ordinal position, but pos-sibly Caloneurodea. It diff ers also from Lusitaneura in the much reduced area of RP.

Caloneurella Carpenter 1934 shares with Lusitaneura the presence of three branches of RP, the posterior branch of MA with small secondary branches emerging from it but it diff ers in the longer ScP ending near apex of RA (Carpenter, 1934).

Pruvostiella Handlirsch 1922 is a caloneurodean genus based on a single species P. lecomtei (Pruvost 1919) based on a rather incomplete wing with several structures missing. Nevertheless, it shares with Lusitaneura n. gen. the apex of ScP near second third of wing length, area of RP relatively broad with 2–3 branches; MA forked. Lusitaneura n. gen. diff ers from Pruvostiella in the absence of small branches of CuPaβ near posterior wing margin.Acknowledgments. Th e authors are grateful to A. Rasnitsyn and O. Béthoux for valuable comments and R. J. Beckemeyer for their availability to provide the fi rst exploratory contacts with some experts in palaeoentomology. Th is work would not have been possible without the generous help of Departamento de Botânica da Faculdade Ciências da Universidade do Porto that provided all the necessary working facilities for microscope photograph of wing. We sincerely thank Dr. Carsten Brauckmann and Dr. Jakub Prokop for their helpful comments on the fi rst version of this paper.

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