Carlos Teixeira developed the study of the

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1 ARTICLE Ann. soc. entomol. Fr. (n.s.), 2010, 46 (1 2) : Lusitaneura covensis n. gen., n. sp., first Caloneurodea from the Carboniferous of Portugal (Insecta: Pterygota: Panorthoptera) 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 Buffon, F Paris, France (4) Departamento & Centro de Geologia da Universidade do Porto, Portugal * Corresponding author 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. Carlos Teixeira developed the study of the Carboniferous insects from Portugal in the years He described alone or in collaboration with the French palaeoentomologist Daniel Laurentiaux numerous fossils, mainly blattodean taxa. In Teixeira (1946), is done the first description of fossil insects from the Upper Stephanian outcrop of São Pedro da Cova (Douro Carboniferous Basin: São Pedro da Cova and Pejão Coalfields), i.e. a Palaeodictyoptera Homaloneura ribeiroi, a Blattinopsidae, and two blattodean Phyloblattidae Phyloblatta fonsecai and P. carringtoni. The 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. Thanks to recent field researches, J. Loureiro and P. Correia discovered new fossil insects in São Pedro da Cova coalfiled outcrops. These are a Palaeodictyoptera, several Blattodeans and the first record of the order Caloneurodea from Portugal, currently described in the present work. This 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 [email protected], [email protected], [email protected], [email protected] Accepté le 28 mai 2009 surprising as this order is already well known from the French classical locality of Commentry similar in age as Douro Basin. Geological setting The 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 confirmed in detail, a model of pullapart 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. The base of the Douro Basin rests unconformable over the Lower Cambrian ( Complexo Xistograuváquico ), and the top is cut by a high-angle reverse 242

2 A first Caloneurodea from the Carniferous of Portugal 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). The Douro Basin filling results from continental intramontane syntectonic sedimentation, with alluvial fans debris flow dominated at the base, and passing upwards to lacustrine/palustrine facies, truncated by a multistory-multichannel fluvial braided system, and the top of the sequence showing lacustrine facies with deltaic features. The 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 megafloristic studies, Lemos de Sousa & Wagner (1983b), put into special evidence the occurrence of Pseudomariopteris cf. busqueti (Zeiller) DanzéCorsin, of cf. Taeniopteris multinervis Weiss and of Neuropteris 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 filiciforme (Von Schlotheim pars) Florin, cf. Lebachia frondosa var. zeilleri Florin and Lebachia parvifolia Florin, and also the presence of Dicranophyllum gallicum Grand Eury and Dicranophyllum lusitanicum (Heer). Figure 1 Main geological regional setting of the Douro basin (lower Stephanian C; NW of Portugal), modified from Pinto de Jesus (2001). 243

3 J. P. Loureiro, P. Correia, A. Nel & A. Pinto de Jesus From palaeozoological studies, Eagar (1983) concludes after the occurrence of Anthraconaia lusitanica (Teixeira) Eagar and of Anthraconaia (?) altissima Eagar. Systematic palaeontology We follow the nomenclature of wing venation and classification 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 coalfield, 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; first 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. Discussion Lusitaneura n. gen. falls in the Caloneurodea Handlirsch, 1937 because it has the diagnostic characters 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 crossveins (visible only in some of them, between RA and RP; absence of anterior branch of RA. The 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 classification within the Caloneurodea, we compare our fossil with the various genera currently attributed to this order (Béthoux et al. 2004). Affinities with Caloneura Brongniart 1885 is unlikely because the apex of ScP is near the second third of wing length and subcostal area is not very broad. Lusitaneura n. gen. differs from Gigagramma Béthoux et al in the absence of any vein distally emerging from CuA+CuPaα, and in the more basal position of apex of ScP. Lusitaneura n. gen. differs from Sthenarocera Brongniart 1885 in the presence of only three posterior branches of RP instead of 8 9. It differs 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) 244

4 A first Caloneurodea from the Carniferous of Portugal more basal position, anterior branch of MA not making a strong curve (also differences with Homaloptila Handlirsch 1919), and RP with three branches, instead of two. Lusitaneura n. gen.differs from Pleisiogramma Carpenter, 1943 in the MA with two branches, instead of being simple, and the apex of ScP in a more basal position. Differences 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. The same last difference concerns Euthygramma Martynov, 1928 and Anomalogramma Carpenter 1943 and Nanogramma Béthoux et al The Gelasopteridae Carpenter 1976 (Gelasopteron Carpenter 1976) are Neoptera of uncertain ordinal position, but possibly Caloneurodea. It differs 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 differs 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. differs from Pruvostiella in the absence of small branches of CuPaβ near posterior wing margin. Acknowledgments. The authors are grateful to A. Rasnitsyn and O. Béthoux for valuable comments and R. J. Beckemeyer for their availability to provide the first exploratory contacts with some experts in palaeoentomology. This 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 first version of this paper. References Béthoux O., Nel A., Lapeyrie J The extinct order Caloneurodea (Insecta: Pterygota: Panorthoptera): wing venation, systematics and phylogenetic relationships. Annales Zoologici 54(2): Carpenter F. M Carboniferous insects from Pennsylvania in the Carnegie Museum and the Museum of Comparative Zoology. Annals of the Carnegie Museum 22: Corsin P., Lemos de Sousa M. J Sur la découverte de deux fossiles dans le Carbonifère de São Pedro da Cova (Portugal). Annales de la Société Géologique du Nord 92: Domingos L. C. F. G., Freire J. L. S., Gomes S. F., Gonçalves F., Pereira E., Ribeiro A The structure of the intramontane Upper Carboniferous basins in Portugal, p in: Lemos de Sousa M. J., Oliveira J. T. (eds). The Carboniferous of Portugal. Memória dos Serviços Geológicos Eagar R. M. C The non-marine bivalve fauna of the Stephanian C of North Portugal, p in: Lemos de Sousa M. J., Oliveira J. T. (eds). The Carboniferous of Portugal. Memória dos Serviços Geológicos Fernandes J. P., Pinto de Jesus A., Teixeira F., Lemos de Sousa M. J Primeiros resultados palinológicos na Bacia Carbonífera do Douro (NO de Portugal) p in: Grandal d Anglade A., Gutiérrez-Marco J. C., Santos Fidalgo L. (eds.). XIII Jornadas de Paleontología Fósiles de Galicia y V Reunión Internacional Proyecto 351 PICG Paleozoico Inferior del Noroeste de Gondwana, A Coruña, 1997, Libro de Resúmenes y Excursiones, Sociedad Española de Paleontología, Madrid. Laurentiaux D., Teixeira C Découverte d un Blattinopsis dans le Carbonifère du Bas-Douro (Portugal). 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5 J. P. Loureiro, P. Correia, A. Nel & A. Pinto de Jesus Wagner R. H The palaeogeographical and age relationships of the Portuguese Carboniferous floras with those of other parts of the Western Iberian Peninsula, p in: Lemos de Sousa M. J., Oliveira J. T. (eds). The Carboniferous of Portugal. Memória dos Serviços Geológicos Wagner R. H., Lemos de Sousa M. J The Carboniferous megafloras of Portugal - A revision of identifications and discussion of stratigraphic ages, p in Lemos de Sousa M. J., Oliveira J. T. (eds). The Carboniferous of Portugal. Memória dos Serviços Geológicos Wagner R. H., Lemos de Sousa M. J Oligocarpia leptophylla (Bunbury), Nomenclatorial history and description of the lectotype. In: Lemos de Sousa M. J., Wagner R. H. (eds.). Papers on the Carboniferous of the Iberian Peninsula (sedimentology, stratigraphy, palaeontology, tectonics and geochronology). Anais da Faculdade de Ciências da Universidade do Porto Suplemento 64: Wagner R. H., Ribeiro A., Lemos de Sousa M. J Bacia Carbonífera do Douro. reinterpretação da geologia do sector Germunde-Choupelo. Recomendações para a investigação geológico-mineira deste sector e de sectores anexos. Fornelo, Relatório inédito, 17 p. 246