With a Review of the Known Chrysopid Fossils (Insecta: Neuroptera)

Advances in Neuropterology. Proceedings of the Third International Symposium on Neuropterology. Berg en Dal. Kmger National Park, R.S.A., 1988. Mansell, M.W. & Aswck H. (Eds). Pretoria, R.S.A. 1990. Pp. 27-32.

Paleochrysopa monteilsensis gen. et sp. nov., a new fossil of Chrysopidae from the Upper Eocene Formation of Monteils (France), with a review of the known chrysopid fossils (Insecta: Neuroptera)

Yves SEMERIA & Andrt NEL Nice, France & La Ciotat, France

RESUME Description d'un nouveau genre et d'une nouvelle espece de Chrysopidae fossile originaire de la rtgion d'A1t.s (Gard, France). I1 s'agit du plus ancien representant connu de la sous-famille des Chrysopinae: Paleochrysopa monteilsensis gen. et sp. nov.

ABSTRACT

The oldest known species of fossil Chrysopinae, from the Monteils Formation, Upper Eocene in the vicinity of Ales (Gard, France), is described as Paleochrysopa monteilsensis gen. et sp. nov. Key words: Neuroptera, Chrysopidae, Chrysopinae, fossil, Paleochrysopa monteilsensis, new genus, new species, Upper Eocene, France.

INTRODUCTION A right forewing of Chrysopidae of Upper Eocene age (Lower Ludian) was recently dis- covered in the lacustrine limestone level of the Monteils Formation, 10 km east of Albs. Although the apex and the base are missing, the wing provides sufficiently distinct characters to warrant its study. Psyllids and psocids are also abundant in the same limestone formation (Nel, unpublished data).

Genus Paleochrysopa gen. nov. Type species: Paleochrysopa monteilsensis sp. nov., described below. Etymology: from the Greek Palaios and Chrysopa: ancient Chrysopa.

Diagnosis: Forewing similar to those of modern genera in the subfamily Chrysopinae, but with two important combined characters, namely, intramedian cell triangular-oval in shape, but very distal to the origin of the radial sector (Rs) and pseudomedian (PsM) not absolutely rectilinear.

Paleochrysopa monteilsensis sp. nov.

Holotype: part and counterpart of a wing fragment, Institut de Paltontologie de MusCum Fig. 1. Diagram of thc Farcwing of Pnleochwsnpa montetciisensis gn. et sp. mv.

Fig. 2. FossiE wing of Puleorh~.~op~rnt~f~rrr!rmsi.\ gen. ct bp. nov In lacus1 r~nt limestone of the Montells Formation. France. A new fossil chrysopid

National Histoire naturelle de Paris, France, IPM-R07719 (and 368 in the Monteils series, collection AndrC Nel). Horizon and type locality: Upper Eocene, Lower Ludian, Monteils Formation, Gard Depart- ment, France. Etymology: derived from the name of the Monteils Formation in which the holotype was found.

Diagnosis: Total length of wing probably 15mm; maximum width 4,9mm. Five cells between the PsM and pseudocubital (PsCu) distal to the intrarnedian cell. Apex of wing rounded. Costal area probably contained 23 to 24 costal veins.

Description: Wing hyaline; length preserved 10mm. The subcosta (Sc) and the radius (R) remain distinct along their entire length, but deviate slightly away from each other near the apex, the maximum deviation between them being 0,215mm. These two veins are covered by setae (very dense in the apical part); other setae, in fewer numbers, can be distinguished on all other veins. Length of the longest seta is 0,18 mm. Apex of wing rounded. The costal area has 16 visible costal veins; costal area narrows progressively towards its apex; maximum width of this area is 0,88mm. The costa (C) and Sc rejoin at 0,98mm from the apex of the wing; there are two small transverse veins between Sc and R close to the end of R and Sc. The PsM is almost rectilinear but less so than in wings of present-day Chrysopinae, and forms an angle of 70" with the inner gradate crossveins. The PsCu is almost rectilinear and parallel to the PsM; it is not preceded by a series of external gradate crossveins; these branch directly on the PsM. Maximum width between the PsM and PsCu is 0,84mm; maximum width between the PsCu and the lower margin of the wing is 0,98mm. The intramedian cell is incomplete; it does not resemble the form of an elongate pentagon. A vein probably existed, which originated at the distal end of this cell and extended in the direction of the PsCu (Fig. 1). The maximum width of the intramedian cell is very distal to the origin of Rs, similar to the more primitive genera of Chrysopidae (Carpenter 1935: 261).

DISCUSSION The examination of this specimen shows features which are not present in Nothochrysi- nae, in fact, the inner gradate crossveins do not extend the PsM at all. It forms, on the contrary, a large open angle with the latter, as is the case in Chrysopinae. On the other hand, it is evident that the intramedian cell, despite its being damaged, has a shape very similar to that found in many modern genera of Chrysopinae (triangular oval cell). Final- ly, the border of this cell is relatively distant from the second transverse vein situated between the Rs and PsM. With few exceptions, the intramedian cell, by its large size in most modern and fossil Nothochrysinae reaches this second vein. In another respect Paleochrysopa manifests two relatively archaic features within the Chrysopinae: the respective positions of the intramedian cell and the origin of Rs almost resembles that of the Nothochrysinae (the modern genus Italochrysa Principi of the subfamily Chrysopinae shows the same feature). The PsM is not absolutely rectilinear but forms a weak "zig-zag". The other fossil of Chrysopinae, Chrysopa sarmatica Handschin, 1937, differs from the new species in the following features: in C, sarmatica the cells of the first row, between R and Rs are nearly of equal height, whilst they are not equal in P. monteilsensis; the wing of the latter is distinctly wider than the former (4,9mm compared to 4mm); the transverse veins which extend from the PsCu to the hind margin of the wing, do not bifurcate in C, sarmatica ; finally, C. sarmatica does not possess the two archaic structures of PsM and respective position of Rs and intramedian cell that is present in Paleochrysopa. Y. Se'me'ria & A. Nel

This discovery completely disrupts our knowledge concerning the fossil Chrysopinae. In fact, the origin of the Chrysopinae may well be in the Upper Cretaceous and, since then, a more ancient separation between the Chrysopinae and the Apochrysopinae will have to be assumed. Yet, the very recent manifestation of the latter subfamily of which we possess no definite fossil, its meagre representation (1 1 genera, 25 species), its absence from Europe and North America (New 1984), rather suggests a relatively recent relation- ship between these two subfamilies. The origin of the Apochrysinae seems to be very ob- scure. Either the Apochrysinae detached from the Chrysopinae during the Tertiary, then the division between the Apochrysinae and the Chrysopinae is paraphyletic, or the Apochry- sinae are very old and their origin is to be found in the Upper Cretaceous, in a common stem group with the Chrysopinae. Paleochrysopa monteilsensis gen. et sp. nov., appears to be a member of the Chrysopinae and presently represents the most ancient known fossil from this group.

COMPLETE LIST OF KNOWN FOSSILS OF CHRYSOPIDAE Mesochrysinae 1. Mesochrysopa zitteli Handlirsch, 1908, Germany, Solnhofen and U. S.S. R., Karatau, Upper Jurassic . 2. Mesypochrysa latipennis Martynov, 1927, U.S. S .R., Karatau, Upper Jurassic. 3. Aristonymphes perfectus Panfilov, 1980, U. S .S.R., Kazakhstan, Upper Jurassic. 4. Macronympha elegans Panfilov, 1980, U. S.S. R., Kazakhstan, Upper Jurassic. 5. Chrysoleonites intactus Panfilov, 1980, U.S.S.R., Kazakhstan, Upper Jurassic. 6. Chrysoleonites plexus Panfilov, 1980, U.S.S.R., Kazakhstan, Upper Jurassic. 7. Nymphoides latus Panfilov, 1980, U.S.S.R., Kazakhstan, Upper Jurassic.

Nothochrysinae 8. Cimbrochrysa moleriensis Schliiter, 1982, Denmark, moler, diatomaceous earth, Lower Eocene. 9. Chrysopidae (undetermined species), Jarzembowski, 1980, England, Isle of Wight, Upper Eocene, Upper Priabonian. 10. Dyspetochrysa vetuscula (Scudder , 1890), U.S.A., Florissant, Lower Oligocene (Wilson 1978). 11. Archaeochrysa fracta (Cockerell, 1914), U.S.A., Florissant, Oligocene. 12. Archaeochrysa paranewis Adams, 1967, U. S.A., Florissant, Oligocene. 13. Palaeochrysa stricta Scudder , 1890, U.S.A., Florissant and Creede Formation, Oligocene. 14. Palaeochrysa wickhami (Cockerell, 1914), U. S .A., Florissant, Oligocene. 15. Palaeochrysa concinnula (Cockerell, 1909), U.S.A., Florissant, Oligocene. 16. Tribochrysa inaequalis Scudder, 1890, U .S.A., Florissant, Oligocene. A new fossil chrysopid

17. Tribochrysa firrnata Scudder, 1890, U. S.A. , Florissant , Oligocene. 18. Notochrysa stampieni Nel & SCmCria, 1986, France, Aix-en-Provence, Upper Oligocene. 19. Archaeochrysa creedei (Carpenter, 1935), U.S.A. Creede Formation, Upper Oligocene or Lower Miocene (Wilson 1978). 20. Notochrysa praeclara Statz, 1936, Germany, Rott, Oligo-Miocene. 21 Chrysopidae (undetermined) designated as Chrysopa sp., Barbu, 1939, Switzerland, Savcel-les-Bains, OltCnie, Miocene (Vindobonian, Sarmatian). 22. Chrysopidae (undetermined species) Martynova, 1949, U.S.S.R., Stavropol, Miocene. 23. Hypochrysodes hercyniensis Schliiter, 1982, Germany, Willershausen, Pliocene. Chryopinae 24. Paleochrysopa monteilsensis SCmCria & Nel, 1990, France, Monteils Formation, Upper Eocene, Lower Ludian. 25. Chrysopa sarmatica Handschin, 1937, Roumania, Mdia Region, Magyar saros, Miocene (Upper Sarmatian or Lower Pontian).

REFERENCES ADAMS, P.A. 1967. A review of the Mesochrysinae and Nothochrysinae (Neuroptera, Chrysopidae). Bulletin of the Museum of Comparative Zoology at Harvard College 135: 214-238. BARBU, I.Z. 1939. Insectes fossiles du tertiaire de l'Olt6nie. Bulletin de la Socie'te Roumaine de Gkologie 4: 119-128. CARPENTER, F.M. 1935. Tertiary insects of the family Chrysopidae. Journal of Paleontology 9: 259-271. CARPENTER, F.M. 1938. Fossil insects from the Creede formation Colorado. Part I: Introduction and order Neuroptera. Psyche 45: 105-109. COCKERELL, T.D. A. 1908. Fossil Chrysopidae. Canadian Entomologist 40: 90-91. COCKERELL, T.D.A. 1909. Two fossil Chrysopidae. Canadian Entomologist 41: 218-219. COCKERELL, T.D.A. 1914. New and little-known insects from the miocene of Florissant, Colorado. Journal of Geology 22: 714-724. HANDLIRSCH, A. 1906-1908. Die fossilen Insekten und die Phylogenie der rezenten formen. Ein Handbuch fir Paliiontologen und Zoologen. 1430 pp. W. Engelmann, Leipzig. HANDSCHIN, A. 1937. Fossile Insekten aus Siebenbiirgen. Mitteilungen der Schweizerichen Entomologischen Gesellschaft 17: 25-29. JARZEMBOWSKI, E.A. 1980. Fossil insects from the Bembridge Marls, Paleogent of the Isle of Wight. Bulletin of the British Museum of Natural History A. Geology 33: 237-293. MARTYNOV, A. 1927. Jurassic fossil insects from Turkestan, 7: some Odonata, Neuroptera, Thysanoptera. Bulletin de I'Acadkmie des Sciences de ['U.R. S. S. 2 1: 757-768. MARTYNOVA, 0. 1949. Mesozoic lacewings (Neuroptera) and their bearing on concepts of phylogeny and systematics of the order. Trudy Paleontologicheskovo Instituta 20: 150-170. NEL, A. and SEMERIA, Y. 1986. Une nouvelle espBce de Chrysopidae fossile du stampien sup6rieur (oligocbne) d'Aix-en-Provence. Neuroptera International 4: 23-30. PANFILOV, D. 1980. New representatives of lacewings (Neuroptera) from the Jurassic of Karatau. In: Dolin, V., Panfilov, D., Ponomarenko, A. & Pritikina, L. (Eds) Mesozoic Fossil Insects. 82-1 11. Naukova Dumka, Kiev. SCHLUTER, T. 1982. Cimbrochrysa moleriensis n.g, n.sp., und Hypochrysa hercyniensis n.sp., zwei fossile Chrysopidae-Arten (Insecta: Planipennia) aus dem europaischen Tertiar. Neues Jarbuch Geol. Paliiont. Mh. 5: 257-264. SCHLUTER, T. 1984. Paleontology and evolutionary relationships. In: Canard, M,, SCmkria, Y. & New, T.R. (Eds) Biology of Chrysopidae. 1-8. Dr W. Junk, The Hague. SCUDDER, S.H. 1890. The fossil insects of North America, with notes on some European species. 11. The Tertiary insects. Report of the U. S. Geological Survey of the Territories 13: 1-734. STATZ, G. 1936. Ueber neue Funde von Neuropteren, Panorpen und Trichopteren aus den Tertiaren Schiefern von Rott am Siebengebirge. Decheniana 93: 208-255. WILSON, M.H.V. 1978. Paleogene insect faunas of western North America, Quaestiones Entomologicae 14: 13-34. Y. Skrneria & A. Nel

Addresses of authors:

Dr Yves SCmCria Dr Andrk Nel Laboratoire de Biologie Animale et de Cytologie 8, avenue Gassion FacultC des Sciences F-13600 La Ciotat Campus Valrose France F-06034 Nice cCdex France Bibliography of the Neuropterida

Bibliography of the Neuropterida Reference number (r#): 7257

Reference Citation: Séméria, Y.; Nel, A. [D.] 1990 [1990.??.??]. Paleochrysopa monteilsensis gen. et sp. nov., a new fossil of Chrysopidae from the Upper Eocene Formation of Monteils (France), with a review of the known chrysopid fossils (Insecta: Neuroptera). Pp. 27-32 in Mansell, M. W.; Aspöck, H. (eds.). Advances in Neuropterology. Proceedings of the Third International Symposium on Neuropterology (3-4 February 1988, Berg en Dal, Kruger National Park, South Africa). South African Department of Agricultural Development, Pretoria. 298 pp.

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