Abstract
A new species of palaeodictyopteridan insect, Heolus martinclosasi sp. nov., based on a partial wing, has been identified in the uppermost Pennsylvanian (Stephanian C local stage) deposits of the Castellar de N’Hug-Camprodon Basin, located in the Surroca-Ogassa coalfield within the Pyrenees of Catalonia, Spain. This species belongs to the Palaeodictyoptera, and is the first of this order described from the Paleozoic deposits of Spain. Heolus martinclosasi sp. nov. is the second species within the relatively small Heolidae family, originally described by Handlirsch in 1906, in the USA. This new species was found alongside remains of in situ fossil plants such as Calamites suckowii, Sphenophyllum oblongifolium, and marattialean tree ferns, all of which grew between river channels. This suggests that these insects possibly lived in lake-like, permanently inundated floodplain deposits.
References
Agnus, A. 1902. Description d’un Neuroptère fossile nouveau. Homoioptera gigantea. Bulletin de la Société Entomologique de France, (N.S.), 7, 259–261. https://doi.org/10.3406/bsef.1902.23126
Álvarez-Ramis, C., Doubinger, J. & Diéguez Jiménez, M.C. (1971) Estudio paleobotánico de la flora de Ogassa (Gerona). Estudios Geológicos, 27, 267–277.
Álvarez-Ramis, C., Pi-Radondy, M. & Doubinger, J. (1979) Sur la flore fossile du Carbonifère de Surroca (Gerona, Espagne). Comptes Rendus de l’Académie des Sciences, Paris, 268, 2559–2561.
Bechly, G. (1996) Morphologische Untersuchungen am Flügelgeäder der rezenten Libellen und deren Stammgruppenvertreter (Insecta; Pterygota; Odonata), unter besonderer Berücksichtigung der Phylogenetischen Systematik und des Grundplanes der Odonata. Petalura, (Special Volume 2), 402 pp.
Bolton, H. (1912) Insect-remains from the Midland and South-Eastern coal measures. Quarterly Journal of the Geological Society, 68, 310–323. https://doi.org/10.1144/GSL.JGS.1912.068.01-04.25
Brongniart, C. (1893) Recherches pour servir à l’histoire des insectes fossiles des temps primaires précédées d’une étude sur la nervation des ailes des insectes. Bulletin de la Société d’Industrie Minérale de Saint-Etienne, 3, 1–491. https://doi.org/10.5962/bhl.title.34754
Carpenter, F.M. (1976) The lower Permian insects of Kansas: part 12. Protorthoptera (continued), Neuroptera, additional Palaeodictyoptera & families of uncertain position. Psyche, 83, 336–376. https://doi.org/10.1155/1976/932123
Carpenter, F.M. (1992) Superclass Hexapoda. In: Moore R.C. & Kaesler, R.L. (Eds), Treatise on Invertebrate Paleontology, Arthropoda 4. The Geological Society of America and the University of Kansas, Boulder, Colorado, (R 3/4), xxii + 655 pp.
Correia, P., Nel, A., Sá, A.A., Domingos, R., Carneiro, A. & Flores, D. (2014) A new Palaeodictyoptera from the Late Carboniferous of Portugal. Annales de la Société Entomologique de France (N.S.), 49, 398–401. https://doi.org/10.1080/00379271.2014.893645
Correia, P., Schubnel, Th. & Nel, A. (2021) What is the roachoid genus Eneriblatta (Dictyoptera: Phyloblattidae) from the Carboniferous of Portugal. Historical Biology, 33, 1–6. https://doi.org/10.1080/08912963.2019.1661407
Gisbert, J. (1986) Els temps tardihercinians. In: P. Santanach (Ed.), Enciclopèdia d’Història Natural dels Països Catalans, Geologia I. Editorial Enciclopèdia Catalana, Barcelona, 197–240.
Goldenberg, F. (1877) Die fossilen Thiere aus der Steinkohlenformation von Saarbrücken. Fauna Saraepontana Fossilis, Mollinger, Saarbrücken, 2, i–iv + 1–54.
Grimaldi, D. & Engel, M.S. (2005) Evolution of the insects. Cambridge University Press, Cambridge, New York, Melbourne, xv + 755 pp.
Guthörl, P. (1930) Neue Insektenfunde aus dem Saarcarbon. Neues Jahrbuch für Mineralogie, Geologie und Paläontologie Abhandlungen, (B), 64, 147–164.
Handlirsch, A. (1904) Les insectes houillers de la Belgique. Mémoires du Musée Royal d’Histoire Naturelle de Belgique, 3, 1–20.
Handlirsch, A. (1906a) (−1908) Die fossilen Insekten und die Phylogenie der rezenten Formen. In: Ein Handbuch für Paläontologen und Zoologen. W. Engelman, Leipzig, p. 1430. https://doi.org/10.5962/bhl.title.34145
Handlirsch, A. (1906b) Revision of American Paleozoic insects. Proceedings of the United States National Museum, 29, 661–820. https://doi.org/10.5479/si.00963801.29-1441.661
Handlirsch, A. (1910) Das erste fossile Insekt aus dem Oberkarbon Westfalens. Verhandlungen der Kaiserlich-Königlichen Zoologisch-Botanischen Gesellschaft in Wien, 60, 249–251.
Handlirsch, A. (1911) New Paleozoic insects from the vicinity of Mazon Creek, Illinois. American Journal of Science, 184, 297–326 + 353–377. https://doi.org/10.2475/ajs.s4-31.185.353
Handlirsch, A. (1919) Revision der paläozoischen Insekten. Denkschriften der Kaiserlichen Akademie der Wissenschaften, Mathematisch-Naturwissenschaftliche Klasse in Wien, 96, 511–592.
Heckel, P.H. & Clayton, G. (2006) The Carboniferous System. Use of the new official names for the subsystems, series, and stages. Geologica Acta, 4, 403–407.
Kukalová, J. (1969a) Revisional study of the order Palaeodictyoptera in the Upper Carboniferous shales of Commentry, France. Part 2. Psyche, 76, 439–486. https://doi.org/10.1155/1969/20732
Kukalová, J. (1969b) Revisional study of the order Palaeodictyoptera in the Upper Carboniferous shales of Commentry, France. Part 1. Psyche, 76, 163–215. https://doi.org/10.1155/1969/74019
Kukalová-Peck, J. (1975) Megasecoptera from the Lower Permian of Moravia. Psyche, 82, 1–19. https://doi.org/10.1155/1975/78215
Kukalová-Peck, J. & Richardson, E.S. Jr. (1983) New Homoiopteridae (Insecta: Paleodictyoptera) with wing articulation from Upper Carboniferous strata of Mazon Creek, Illinois. Canadian Journal of Zoology, 61, 1670–1687. https://doi.org/10.1139/z83-218
Laurentiaux, D. (1950) Les insectes houillers du Limbourg hollandais. Mededelingen van de Geologische Stichling, 4, 13–22.
Laurentiaux, D. & Laurentiaux-Vieira, F. (1951) Observations sur les Paléodictyoptères Breyeriidae. Bulletin de la Société Géologique de France, (6), 1 (7), 585–596. https://doi.org/10.2113/gssgfbull.S6-I.7.585
Laurentiaux-Vieira, F. & Laurentiaux, D. (1986) Paléodictyoptère nouveau du Namurien belge. Annales de la Société Géologique du Nord, 105, 187–193.
Logghe, A., Garrouste, R., Steyer, J.-S., Pouillon, J.-M. & Nel, A. (2022) The first Palaeodictyoptera (Insecta) from the Carboniferous-Permian basin of Graissessac (France). Historical Biology, 34, 486–492. https://doi.org/10.1080/08912963.2021.1929953
Martín-Closas, C. & Martínez-Roig, D. (2007) Plant taphonomy and palaeoecology of Stephanian limnic wetlands in the eastern Pyrenees (Catalonia, Spain) Comptes Rendus Palevol, 6, 437–449. https://doi.org/10.1016/j.crpv.2007.09.007
Martynov, A.V. (1931) On some remarkable Odonata from the Permian of Archangelsk district. Izvestiya Akademii Nauk SSSR, 1931 (1), 141–147.
Meunier, F. (1908) Nouveaux Mégasécoptèrides et un nouveau Paléodictyoptère de Commentry. Bulletin du Museum National d’Histoire Naturelle, 14, 172–175.
Nel, A., Santos, A.A., Hernández-Orúe, A., Wappler, T., Diez, J.B. & Peñalver, E. (2022) The first representative of the roachoid family Spiloblattinidae (Insecta, Dictyoptera) from the Late Pennsylvanian of the Iberian Peninsula. Insects, 13, 828. https://doi.org/10.3390/insects13090828
Özdikmen, H. (2008) Some nomenclatural changes for Blattodea and Dictyoneurida (= Palaeodictyoptera). Munis Entomology & Zoology, 3, 745–748.
Peñalver, E., Martínez-Delclòs, X. & Arillo, A. (1999) Yacimientos con insectos fósiles en España. Revista Española de Paleontología, 14, 231–245. https://doi.org/10.7203/sjp.23796
Prokop, J. & Engel, M.S. (2019) Palaeodictyopterida. Current Biology, 29, 306–309. https://doi.org/10.1016/j.cub.2019.02.056
Prokop, J. & Nel, A. (2004) A new genus and species of Homoiopteridae from the Upper Carboniferous of the Intra-Sudetic Basin, Czech Republic (Insecta, Palaeodictyoptera). European Journal of Entomology, 101, 583–589. https://doi.org/10.14411/eje.2004.082
Prokop, J. & Ren, D. (2007) New significant fossil insects from the Upper Carboniferous of Ningxia in northern China (Palaeodictyoptera, Archaeorthoptera) European Journal of Entomology, 104, 267–275. https://doi.org/10.14411/eje.2007.041
Prokop, J., Nel, A. & Tenny, A. (2010) On the phylogenetic position of the palaeopteran Syntonopteroidea (Insecta: Ephemeroptera), with a new species from the Upper Carboniferous of England. Organisms Diversity & Evolution, 10, 331–340. https://doi.org/10.1007/s13127-010-0022-2
Prokop, J., Tippeltova, Z., Roques, P. & Nel, A. (2013) A new genus and species of Breyeriidae and wings of immature stages from the Upper Carboniferous, Nord Pas-de-Calais, France (Insecta: Palaeodictyoptera). Insect Systematics & Evolution, 44, 117–128. https://doi.org/10.1163/1876312X-44032098
Riek, E.F. (1976) Neosecoptera, a new insect suborder based on specimen discovered in the late Carboniferous of Tasmania. Alcheringa, 1, 227–234. https://doi.org/10.1080/03115517608619072
Scudder, S.H. (1885) Systematische Übersicht der fossilen Myriapoden [sic], Arachnoideen, und Insekten. Pp. 721–831. In: Zittel, K.A. (Ed.), Handbuch der Palaeontologie. Abteilung I, Zweiter Band, Palaeozoologie. Oldenbuch, Munich, 831 pp. https://doi.org/10.5962/bhl.title.2492
Scudder, S.H. (1893) Insect fauna of the Rhode Island coal field. Bulletin of the United States Geological Survey of the Territories, 101, 1–21.
Sinitshenkova, N.D. (1992) Two new insect species (Insecta: Dictyoneurida = Palaeodictyoptera, Perlida = Plecoptera) from the Late Permian of South Mongolia. Trudy Sovmestnaya Sovetsko-Mongol’skaya Paleontologicheskaya Ekspeditsiya, 4, 98–100.
Sinitshenkova, N.D. (2002) Chapter 2.2.1.2.3. Superorder Dictyoneurida Handlirsch, 1906 (= Palaeodictyopteroidea) pp. 115–124. In: Rasnitsyn, A.P. & Quicke, D.L.J. (Eds), History of insects. Kluwer Academic Publishers, Dordrecht (Boston, London), pp. xi + 517.
Sroka, P., Staniczek, A.H. & Bechly, G. (2015) Revision of the giant pterygote insect Bojophlebia prokopi Kukalová-Peck, 1985 (Hydropalaeoptera: Bojophlebiidae) from the Carboniferous of the Czech Republic, with the first cladistic analysis of fossil palaeopterous insects. Journal of Systematic Palaeontology, 13, 963–982. https://doi.org/10.1080/14772019.2014.987958
Tosal, A., Pàmies, J. & Martín-Closas, C. (2022) Plant taphonomy and palaeoecology of Pennsylvanian wetlands from the Erillcastell Basin of the eastern Pyrenees, Catalonia, Spain. Palaeogeography, Palaeoclimatology, Palaeoecology, 605, 111234. https://doi.org/10.1016/j.palaeo.2022.111234
Wagner, R.H. (2004) Gondomaria grandeuryi (Zeiller) Wagner & Castro, 1998, in the context of an upper Stephanian flora from Surroca (prov. Girona, Catalonia, Spain). Treballs del Museu de Geologia de Barcelona, 12, 53–67.
Wagner, R.H. & Álvarez-Vázquez, C. (2010) The Carboniferous floras of the Iberian Peninsula: A synthesis with geological connotations. Review of Palaeobotany and Palynology, 162, 239–324. https://doi.org/10.1016/j.revpalbo.2010.06.005