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Type: Article
Published: 2022-05-05
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A revision of Chrysopidae (Neuroptera) from the late Eocene Florissant Formation, Colorado, with description of new species

Federal Scientific Center of the East Asia Terrestrial Biodiversity, Far East Branch of the Russian Academy of Sciences, Vladivostok, 960022, Russia
Department of Earth Sciences, University of Oxford, Oxford OX1 3AN, United Kingdom
Department of Biological Sciences, Simon Fraser University, 8888 University Drive, Burnaby, British Columbia, V5A 1S6, Canada . Museum of Comparative Zoology, 26 Oxford Street, Cambridge, Massachusetts, 02138, United States of America 5Royal British Columbia Museum, 675 Belleville Street, Victoria, British Columbia, V8W 9W2, Canada
Neuroptera Chrysopidae Nothochrysinae Florissant Formation late Eocene


The green lacewings (Neuroptera: Chrysopidae) of the late Eocene Florissant Formation (Colorado, USA) are revised. Thirteen species in six genera of Nothochrysinae are recognized, including four new species: Archaeochrysa cockerelli sp. nov., Cimbrochrysa americana sp. nov., C. major sp. nov., and Lithochrysa meyeri sp. nov. The validity of the genus Dyspetochrysa Adams, 1967 is tenuous; it might be a synonym of Archaeochrysa Adams, 1967. Lithochrysa Carpenter, 1935 and L. ferruginea (Cockerell, 1909) are considered a valid genus and species. A lectotype and paralectotype of Tribochrysa firmata Scudder, 1890 are designated. The genus Cimbrochrysa Schlüter, 1982 is recorded from North America for the first time. Eocene chrysopid assemblages are briefly analyzed. The late Eocene Florissant assemblage is less diverse than those of early Eocene North America, and greatly differs from late Eocene European assemblages.



  1. Adams, P.A. (1957) A new genus and new species of Chrysopidae from the western United States, with remarks on the wing venation of the family (Neuroptera). Psyche, 63, 67–74. [for 1976]
    Adams, P.A. (1967) A review of the Mesochrysinae and Nothochrysinae (Neuroptera: Chrysopidae). Bulletin of the Museum of Comparative Zoology, 135, 215–238.
    Adams, P.A. & Penny, N.D. (1992a) Review of the South American genera of Nothochrysinae (Insecta: Neuroptera: Chrysopidae). In: Canard, M., Aspöck, H. & Mansell, M.W. (Eds.), Current Research in Neuropterology. Proceedings of the Fourth International Symposium on Neuropterology, 24–27 June 1991, Bagnères-de-Luchon, Haute-Garonne, France. Privetely printed, Toulouse, pp. 35–41.
    Adams, P.A. & Penny, N.D. (1992b) New genera of Nothochrysinae from South America (Neuroptera: Chrysopidae). Pan-Pacific Entomologist, 68, 216–221.
    Allen, S.E., Lowe, A.J., Peppe, D.J. & Meyer, H.W. (2020) Paleoclimate and paleoecology of the latest Eocene Florissant flora of central Colorado, U.S.A. Palaeogeography, Palaeoclimatology, Palaeoecology, 551, 109678.
    Archibald, S.B. & Cannings, R.A. (2019) Fossil dragonflies (Odonata: Anisoptera) from the early Eocene Okanagan Highlands, western North America. The Canadian Entomologist, 151, 783–816.
    Archibald, S.B., Johnson, K.R., Mathewes, R.W. & Greenwood, D.R. (2011) Intercontinental dispersal of giant thermophilic ants across the Arctic during early Eocene hyperthermals. Proceedings of the Royal Society B, 278, 3679–3686.
    Archibald, S.B. & Makarkin, V.N. (2006) Tertiary giant lacewings (Neuroptera: Polystoechotidae): revision and description of new taxa from western North America and Denmark. Journal of Systematic Palaeontology, 4, 119–155 + 307 (errata).
    Archibald, S.B. & Makarkin, V.N. (2015) A new species of Archaeochrysa Adams (Neuroptera: Chrysopidae) from the Early Eocene of Driftwood Canyon, British Columbia, Canada. Canadian Entomologist, 147, 359–369.
    Archibald, S.B. & Makarkin, V.N. (2017) A new fossil green lacewing (Neuroptera: Chrysopidae) from the early Eocene Driftwood Canyon, Canada. Zootaxa, 4324 (2), 397–400.
    Archibald, S.B. & Makarkin, V.N. (2021) Early Eocene snakeflies (Raphidioptera) of western North America from the Okanagan Highlands and Green River Formation. Zootaxa, 4951(1), 41–79.
    Archibald, S.B., Makarkin, V.N., Greenwood, D.R. & Gunnell, G.F. (2014) The Red Queen and Court Jester in green lacewing evolution: bat predation and global climate change. Palaios, 29, 185–191.
    Bather, F.A. (1909) Visit to the Florissant exhibit in the British Museum (Natural History). Proceedings of the Geologists Association of London, 21, 159–165.
    Breitkreuz, L., Winterton, S. & Engel, M.S. (2017) Wing tracheation in Chrysopidae and other Neuropterida (Insecta): A resolution of the confusion about vein fusion. American Museum Novitates, 3890, 1–44.
    Carpenter, F.M. (1935) Tertiary insects of the family Chrysopidae. Journal of Paleontology, 9, 259–271.
    Carpenter, F.M. (1943) Osmylidae of the Florissant shales, Colorado (Insecta-Neuroptera). American Journal of Science, 241, 753–760.
    Carpenter, F.M. (1960) Fossil Nemopteridae (Neuroptera). Psyche, 66(1/2), 20–24.
    Carpenter, F.M. (1992) Superclass Hexapoda. In: Kaesler, R.L. (Ed.), Treatise on Invertebrate Paleontology. Part R. Arthropoda 4. Vols. 3 & 4. Geological Society of America, Boulder, Colorado & University of Kansas, Lawrence, Kansas, pp. 1–655.
    Cockerell, T.D.A. (1907a) Some Old-World types of insects in the Miocene of Colorado. Science, 26, 446–447.
    Cockerell, T.D.A. (1907b) An enumeration of the localities in the Florissant Basin, from which fossils were obtained in 1906. Bulletin of the American Museum of Natural History, 23, 127–132.
    Cockerell, T.D.A. (1908a) Fossil Chrysopidae. The Canadian Entomologist, 40, 90–91.
    Cockerell, T.D.A. (1908b) Fossil Osmylidae (Neuroptera) in America. The Canadian Entomologist, 40, 341–342.
    Cockerell, T.D.A. (1909a) A catalogue of the generic names based on American insects and arachnids from the tertiary rocks, with indications of the type species. Bulletin of the American Museum of Natural History, 26, 77–86.
    Cockerell, T.D.A. (1909b) Two fossil Chrysopidae. The Canadian Entomologist, 41, 218–219.
    Cockerell, T.D.A. (1913) Some fossil insects from Florissant, Colorado. Proceedings of the United States National Museum, 44, 341–346.
    Cockerell, T.D.A. (1914) New and little-known insects from the Miocene of Florissant, Colorado. Journal of Geology, 22, 714–724.
    Esben-Petersen, P. (1917) Australian Neuroptera. Part iii. Proceedings of the Linnean Society of New South Wales, 42, 203–219.
    Evanoff, E., McIntosh, W.C. & Murphey, P.C. (2001) Stratigraphic summary and 40Ar/39Ar geochronology of the Florissant Formation, Colorado. In: Evanoff, E., Gregory-Wodzicki, K.M. & Johnson, K.R. (Eds.), Fossil flora and stratigraphy of the Florissant Formation, Colorado. Proceedings of the Denver Museum of Nature & Science, Series 4, No. 1, pp. 1–16.
    Hagen, H.A. (1866) Hemerobidarum Synopsis synonymica. Stettiner Entomologische Zeitung, 27, 369–462.
    Handlirsch, A. (1906–1908) Die fossilen Insekten und die Phylogenie der rezenten Formen. Ein Handbuch für Paläontologen und Zoologen. W. Engelmann, Leipzig, ix+1430 pp., 51 pls. [Issued in 1906 (pp. 1–640), 1907 (pp. 641–1120), 1908 (pp. 1120–1430)].
    Handlirsch, A. (1913) Subphylum C. Insecta (Hexapoda). Insects. In: Zittel, K.A. von., Text-book of paleontology, edited by Charles R. Eastman. Vol. 1. 2nd Edition. Macmillan and Co., London, pp. 794–822.
    Handlirsch, A. (1920–1921) Siebentes Kapitel. Palaeontologie. In: Schröder, C. (Ed.), Handbuch der Entomologie. 3. Band. Gustav Fischer, Jena, pp. 117–306. [issued in 1920 (Lieferung 5, pp. 113–208), 1921 (Lieferungs 6 and 7 (part), pp. 209–306). Vol. 3 was published as a separate book in 1925]
    Huang, D.Y., Lian, X.N., Jouault, C., Gao, J. & Nel, A. (2021) First green lacewing (Insecta: Neuroptera: Chrysopidae) from the Paleocene Sanshui Basin of Guangdong, South China. Historical Biology. [in press]
    Jarzembowski, E.A. (1980) Fossil insects from the Bembridge Marls, Palaeogene of the Isle of Wight. Bulletin of the British Museum (Natural History), Geology, 33, 237–293.
    Khramov, A. (2018) A new assemblage of Early Cretaceous green lacewings (Chrysopidae: Neuroptera) from Transbaikalia. In: Ross, A., McGowan, A. (Eds), Earth and Environmental Science Transactions of the Royal Society of Edinburgh, 107 (2/3), 195–202. [for 2017]
    Khramov, A.V., Liu, Q., Zhang, H.C. & Jarzembowski, E.A. (2016) Early green lacewings (Insecta: Neuroptera: Chrysopidae) from the Jurassic of China and Kazakhstan. Papers in Palaeontology, 2, 25–39.
    Kimmins, D.E. (1952). Some new Australian Chrysopidae. Annals and Magazine of Natural History, Series 12, 5, 69–81.
    Lahmers, K. (2011) A primer on lacewings and a quick look at a prehistoric lacewing captured in Florissant’s prehistoric lake shales. Pick & Pack. The Bulletin of Colorado Springs Mineralogical Society, 51 (3), 6–8.
    Laurentiaux, D. (1953) Сlasse des Insectes (Insecta Linné, 1758). In: Piveteau, J. (Ed.), Traité de Paléontologie. Vol. 3. Masson et Cie, Paris, pp. 397–527.
    Linnaeus, C. (1758) Systema naturae per regna tria naturae secundum classes, ordines, genera, species, cum characteribus, differentiis, synonymis, locis. Vol. 1. 10th Edition. Salvii, Holmiae, 824 pp.
    Lu, X.M., Wang, B., Ohl, M. & Liu, X.Y. (2018) The first green lacewing (Insecta: Neuroptera: Chrysopidae) from the mid-Cretaceous amber of Myanmar. Zootaxa, 4399 (4), 563–570.
    Makarkin, V.N. (1991) Miocene Neuroptera from North Caucasus and Sikhote-Alin. Paleontologicheskii Zhurnal, 1991(1), 57–68 (In Russian, English summary).
    Makarkin, V.N. (1998) New Tertiary Neuroptera from the Russian Far East. Tertiary Research, 18 (3/4), 77–83.
    Makarkin V.N. (2014) A new fossil green lacewing (Neuroptera: Chrysopidae) from the Eocene Tadushi Formation, eastern Sikhote-Alin. Far Eastern Entomologist, 272, 1–7.
    Makarkin, V.N. (2017) A remarkable new genus of Protosmylinae (Neuroptera: Osmylidae) from late Eocene Florissant, Colorado. Zootaxa, 4268 (4), 581–587.
    Makarkin, V.N. & Archibald, S.B. (2013) A diverse new assemblage of green lacewings (Insecta: Neuroptera: Chrysopidae) from the Early Eocene Okanagan Highlands, western North America. Journal of Paleontology, 87, 123–146.
    Makarkin, V.N., Archibald, S.B., Mathewes, R.W. (2021) New Protosmylinae (Neuroptera: Osmylidae) from the early Eocene of western North America, with taxonomic remarks. Zootaxa, 4980 (1), 142–156.
    Makarkin, V.N., Wedmann, S. & Weiterschan, T. (2016) A new genus of Hemerobiidae (Neuroptera) from Baltic amber, with a critical review of the Cenozoic Megalomus-like taxa and remarks on the wing venation variability of the family. Zootaxa, 4179 (3), 345–370.
    Makarkin, V.N., Wedmann, S. & Weiterschan, T. (2018) The first green lacewings from the late Eocene Baltic amber. Acta Palaeontologica Polonica, 63, 527–537.
    Martins-Neto, R.G. & Vulcano, M.A. (1989) Neurópteros (Insecta: Planipennia) da Formação Santana (Cretáceo Inferior), Bacia do Araripe, Nordeste do Brasil. I—Família Chrysopidae. Anais da Academia Brasileira de Ciencias, 60, 189–201. [for 1988]
    McLachlan, R. (1868) A monograph of the British Neuroptera-Planipennia. Transactions of the [Royal] Entomological Society of London, 16, 145–224.
    Meyer, H.W. (2003) The Fossils of Florissant. Smithsonian Books, Washington, 258 pp.
    Navás, L. (1910) Crisópidos (Ins. Neur.) nuevos. Brotéria, Zoológica, 9, 38–59.
    Nel, A., Delclòs, X. & Hutin, A. (2005) Mesozoic chrysopid-like Planippenia: a phylogenetic approach (Insecta: Neuroptera). Annales de la Société Entomologique de France, 41, 29–68.
    Nel, A. & Jarzembowski, E. (2019) New lacewings from the Insect Bed (late Eocene) of the Isle of Wight (Neuroptera: Nemopteridae, Chrysopidae, Hemerobiidae). Earth and Environmental Science Transactions of the Royal Society of Edinburgh, 110 (3/4), 397–403.
    Ngô-Muller, V., Blot, K. & Nel, A. (2019) A new species of Nothochrysa from the latest Oligocene of France (Insecta: Neuroptera: Chrysopidae). Palaeoentomology, 2 (2), 111–114.
    O’Brien, N.R., Meyer, H.W. & Harding, I.C. (2008) The role of biofilms in fossil preservation, Florissant Formation, Colorado. In: Meyer, H.M. & Smith, D.M. (Eds.), Paleontology of the Upper Eocene Florissant Formation, Colorado. Geological Society of America Special Paper, 435, 159–177.
    Oswald, J.D. (1993) Revision and cladistic analysis of the world genera of the family Hemerobiidae (Insecta: Neuroptera). Journal of New York Entomological Society, 101, 143–299.
    Oswald, J.D. (2022) Bibliography of the Neuropterida. Lacewing Digital Library. Research Publication No. 2. Available from: (accessed 7 February 2022)
    Peñalver, E., Nel, A. & Martínex-Delclòs, X. (1995) New Nothochrysinae from the spanish Miocene (Neuroptera, Chrysopidae). Bulletin de la Société Entomologique de France, 100, 481–487.
    Penney, D. & Jepson J.E. (2014) Fossil insects. An Introduction to Palaeontology. Siri Scientific Press, Manchester, 224 pp. [with artwork by Richard Bizley]
    Petrulevičius, J.F., Nel, A., Rust, J., Bechly, G. & Kohls, D. (2007) New Paleogene Epallagidae (Insecta: Odonata) recorded in North America and Europe. Biogeographic implications. Alavesia, 1, 15–25.
    Popov, V.K. & Grebennikov, A.V. (2001) New data on the age of effusives from the Bogopol Formation in Primorye. Tikhookeanskaya Geologiya, 3, 47–54. [in Russian]
    Ren, D. & Guo, Z.G. (1996) On the new fossil genera and species of Neuroptera (Insecta) from the Late Jurassic of northeast China. Acta Zootaxonomica Sinica, 21, 461–479.
    Rodeck, H.G. (1938) Type specimens of fossils in the University of Colorado Museum. The University of Colorado Studies, Series A, 25, 281–304.
    Schlüter, T. (1982) Cimbrochrysa moleriensis n. g. n. sp. und Hypochrysa hercyniensis n. sp., zwei fossile Chrysopidae-Arten (Insecta: Planipennia) aus dem europäischen Tertiär. Neues Jahrbuch für Geologie und Paläontologie, Monatshefte, 5, 257–264.
    Schlüler, T. (1984) Phylogeny of Chrysopidae. In: Canard, M., Séméria, Y. & New, T.R. (Eds.), Biology of Chrysopidae. Dr. W. Junk Publishers, the Hague, pp. 1–8.
    Schneider, W.G. (1851) Symbolae ad monographiam generis Chrysopae, Leach. Hirt, Vratislaviae, 178 pp.
    Scudder, S.H. (1882) Notes on some of the Tertiary Neuroptera of Florissant, Colo., and Green River, Wyoming Terr. Proceedings of the Boston Society of Natural History, 21, 407–409.
    Scudder, S.H. (1883) The tertiary Lake Basin at Florissant, Colo., between South and Hayden Parks. Annual Report of the United States Geological and Geographical Survey of the Territories, 12, 271–293.
    Scudder, S.H. (1885) Systematische Übersicht der fossilen Myriopoden, Arachnoideen und Insekten. In: Zittel, K.A., Handbuch der Palaeontologie. Abt. 1. Palaeozoologie. Bd 2. Mollusca und Arthropoda. Verlag von R. Oldenbourg, München, Leipzig, pp. 721–831.
    Scudder, S.H. (1890) The Tertiary insects of North America. Report of the United States Geological Survey of the Territories, 13, 1–734, 28 pls.
    Scudder, S.H. (1891) Index to the known fossil insects of the world including myriapods and arachnids. Bulletin of the United States Geological Survey, 71, 1–744.
    Simonsen, T.J., Archibald, S.B., Rasmussen, J.A., Sylvestersen, R.L., Olsen, K. & Ware, J.L. (2022) Danowhetaksa gen. nov. with two species from the early Eocene Ølst Formation from Denmark, the first Palearctic Whetwhetaksidae (Odonata: Cephalozygoptera). Zootaxa (in press).
    Séméria, Y. & Nel, A. (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). In: Mansell, M.W. & Aspöck, H. (Eds.), Advances in Neuropterology. Proceedings of the Third International Symposium on Neuropterology (Berg in Dal, Kruger National Park, 1988). South African Department of Agricultural Development, Pretoria, pp. 27–32.
    Tauber, C.A. (2014) Nothochrysinae (Neuroptera: Chrysopidae): New larval description and generic synonymy, with a consideration of generic relationships. Psyche, ID 839261, 1–10.
    Tauber, C.A. (2019a) South American Nothochrysinae (Neuroptera, Chrysopidae): I. Description of Nothochrysa ehrenbergi sp. nov. ZooKeys, 866, 1–18. []
    Tauber, C.A. (2019b) South American Nothochrysinae (Neuroptera, Chrysopidae): II. Redescription of Leptochrysa prisca Adams & Penny. ZooKeys, 866, 19–38. []
    Tjeder, B. (1966) Neuroptera-Planipennia. The Lace-wings of Southern Africa. 5. Family Chrysopidae. In: Hanström, B., Brinck, P. & Rudebec, G. (Eds.), South African Animal Life. Vol. 12. Swedish Natural Science Research Council, Stockholm, pp. 228–534.
    Willmann, R. (1993) Insekten aus der Fur-Formation von Dänemark (Moler, ob. Paleozän / unt. Eozän?). 8. Zwei neue Vertreter der Chrysopidae (Neuroptera). Neues Jahrbuch für Geologie und Paläontologie, Monatshefte, 4, 239–245.
    Willmann, R. & Brooks, S.J. (1991) Insekten aus der Fur-Formation von Dänemark (Moler, ob. Paleozän/ unt. Eozän?). 6. Chrysopidae (Neuroptera). Meyniana, 43, 125–135.
    Winterton, A.L., Gillung, J.P., Garzón-Orduña, I.J., Badano, D., Breitkreuz, L.C.V., Duelli, P.R., Engel, M.S., Liu, X.Y., Machado, R.J P., Mansell, M., Mochizuki, A., Penny, N.D., Tauber, C.A. & Oswald, J.D. (2019) Evolution of green lacewings (Neuroptera: Chrysopidae): an anchored phylogenomics approach. Systematic Entomology, 44, 514–526.
    Yang, C.K. & Hong, Y.C. (1990) Drakochrysa, an Early Cretaceous new genus of Chrysopidae (Insecta: Neuroptera) from Laiyang Basin, Shandong Province. Geoscience, 4 (4), 15–26, 1 pl.
    Zaborac-Reed, S.J. & Leopold, E.B. (2016) Determining the paleoclimate and elevation of the late Eocene Florissant flora: support the coexistence approach. Canadian Journal of Earth Sciences, 53, 565–573.
    Zhang, T.W., Luo, C.S., Shi, C.F., Yang, Q. & Ren, D. (2020) New species of green lacewings (Insecta, Neuroptera) from the Early Cretaceous of China. Cretaceous Research, 115, 104564.