Abstract
The neuropterid (Neuroptera and Raphidioptera) fauna of the middle Eocene Coal Creek Member (Kishenehn Formation), U.S.A. is documented. Three families of Neuroptera (Ascalaphidae, Chrysopidae, Hemerobiidae) and two families of Raphidioptera (Raphidiidae, Inocelliidae) are recorded. Five new species and three new genera are described: Pseudoameropterus ambiguus gen. et sp. nov. (Ascalaphidae), Minimochrysa latialata gen. et sp. nov., Palaeochrysa greenwalti sp. nov., P. minor sp. nov. (Chrysopidae: Nothochrysinae), Macrostigmoraphia diluta gen. et sp. nov. (Raphidiidae). Two indeterminate species are also recorded: Megalomus-group gen. et sp. indet. (Hemerobiidae) and Neuroptera fam. gen. et sp. indet. The only previously described neuropterid from the deposit is the raphidiopteran Paraksenocellia australis Makarkin et al. 2019 (Inocelliidae). The neuropterid assemblage suggests a subtropical semi-arid climate of the Coal Creek Member.
References
- Adams, P.A. (1970) A review of the New World Dilaridae. Postilla, 148, 1–30.
- Ardila-Camacho, A., Noriega, J.A. & Acevedo-Ramos, F. (2019) New genera records of split-eyed owlflies (Neuroptera: Myrmeleontidae: Ascalaphinae) from Colombia. Papéis Avulsos de Zoologia, 59, e20195951. http://doi.org/10.11606/1807-0205/2019.59.51
- 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 (4), 359–369. https://doi.org/10.4039/tce.2014.53
- 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. https://doi.org/10.11646/zootaxa.4324.2.13
- 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. [erratum: 4964 (3), 600] https://doi.org/10.11646/zootaxa.4951.1.2
- 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 (5), 185–191. https://doi.org/10.2110/palo.2013.089
- Aspöck, U. & Aspöck, H. (2004) Two significant new snakeflies from Baltic amber, with discussion on autapomorphies of the order and its included taxa (Raphidioptera). Systematic Entomology, 29, 11–19. https://doi.org/10.1111/j.1365-3113.2004.00245.x
- Aspöck, U. & Aspöck, H. (2008) Phylogenetic relevance of the genital sclerites of Neuropterida (Insecta: Holometabola). Systematic Entomology, 33, 97–127. https://doi.org/10.1111/j.1365-3113.2007.00396.x
- Aspöck, H. & Aspöck, U. (2014) Die nördlichen und südlichen Verbreitungsgrenzen der Ordnung Raphidioptera (Insecta: Endopterygota: Neuropterida). DGaaE-Nachrichten, 28, 16–20.
- Aspöck, U., Aspöck, H. & Rausch, H. (1992) Rezente Südgrenzen der Ordnung Raphidioptera in Amerika (Insecta: Neuropteroidea). Entomologia Generalis, 17, 169–184. https://doi.org/10.1127/entom.gen/17/1992/169
- Aspöck, U., Liu, X.Y., Rausch, H. & Aspöck, H. (2011) The Inocelliidae of southeast Asia: a review of present knowledge (Raphidioptera). Deutsche Entomologische Zeitschrift, Neue Folge, 58, 259–274. https://doi.org/10.1002/mmnd.201100029
- Aspöck, U., Plant, J.D. & Nemeschkal, H.L. (2001) Cladistic analysis of Neuroptera and their systematic position within the Neuropterida (Insecta: Holometabola: Neuropterida: Neuroptera). Systematic Entomology, 26, 73–86. https://doi.org/10.1046/j.1365-3113.2001.00136.x
- Breitkreuz, L.C.V., Winterton, S.L. & 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. https://doi.org/10.1206/3890.1
- Brooks, S.J. & Barnard, P.C. (1990) The green lacewings of the world: a generic review (Neuroptera: Chrysopidae). Bulletin of the British Museum of Natural History, Entomology, 59, 117–286.
- Cai, C.Y., Tihelka, E., Liu, X.Y. & Engel, M.S. (2023) Improved modelling of compositional heterogeneity reconciles phylogenomic conflicts among lacewings. Palaeoentomology, 6 (1), 49–57. https://doi.org/10.11646/palaeoentomology.6.1.8
- Constenius, K.N. (1996) Late Paleogene extensional collapse of the Cordilleran foreland fold and thrust belt. Geological Society of America Bulletin, 108, 20–39.
- Constenius, K.N., Dawson, M.R., Pierce, H.G., Walter, R.C. & Wilson, M.V.H. (1989) Reconnaissance paleontologic study of the Kishenehn Formation, northwestern Montana and southeastern British Columbia. In: French, D.E. & Grabb, R.F. (Eds.), 1989 Field Conference Guidebook: Montana Centennial Edition. Geological Resources of Montana. Vol. 1. Montana Geological Society, Billings, Motana, pp. 189–203.
- Downen, M.R. & Selden, P.A. (2020) Fossil spiders (Araneae) from the Eocene Kishenehn Formation of Montana, USA. Palaeontologia Electronica, 23 (3), a56. https://doi.org/10.26879/1135
- Dawson, M.R. & Constenius, K.N. (2018) Mammalian fauna of the Middle Eocene Kishenehn Formation, Middle Fork of the Flathead River, Montana. Annals of Carnegie Museum, 85 (1), 25–60. https://doi.org/10.2992/007.085.0103.
- Esben-Petersen, P. (1922) New species of Neuroptera in the British Museum. Annals and Magazine of Natural History, Series 9, 10, 617–621.
- Greenwalt, D.E. & Labandeira, C. (2013) The Amazing fossil insects of the Eocene Kishenehn Formation in Northwestern Montana. Rocks & Minerals, 88, 434–441. https://doi.org/10.1080/00357529.2013.809972
- Greenwalt, D.E., Rose, T.R., Siljestrom, S.M., Goreva, Y.S., Constenius, K.N. & Wingerath, J.G. (2015) Taphonomy of the fossil insects of the middle Eocene Kishenehn Formation. Acta Palaeontologica Polonica, 60, 931–947. http://doi.org/10.4202/app.00071.2014
- Gerstaecker, [C.E.] A. (1885) Vier Decaden von Neuropteren aus der Familie Megaloptera Burm. Mittheilungen aus dem Naturwissenschaftlichen Verein für Neu-Vorpommern und Rugen, 16 , 1–49. [for 1884]
- Gerstaecker, [C.E.] A. (1888) Weitere Beiträge zur artenkenntniss der Neuroptera Megaloptera. Mittheilungen aus dem Naturwissenschaftlichen Verein für Neu-Vorpommern und Rugen, 19, 89–130. [for 1887]
- Groves, C. & Shekelle, M. (2010) The genera and species of Tarsiidae. International Journal of Primatology, 31, 1071–1082. https://doi.org/10.1007/s10764-010-9443-1
- Harbach, R.E. & Greenwalt, D. (2012) Two Eocene species of Culiseta (Diptera: Culicidae) from the Kishenehn Formation in Montana. Zootaxa, 3530, 25–34.
- Jones, J.R. (2019) Total-evidence phylogeny of the owlflies (Neuroptera, Ascalaphidae) supports a new higher-level classification. Zoologica Scripta, 48 (6), 761–782. https://doi.org/10.1111/zsc.12382
- Latreille, P.A. (1802) Histoire naturelle, générale et particulière de Crustacés et des Insectes. Vol. 3. Dufart, Paris, 467 pp.
- Latreille, P.A. (1810) Considérations générales sur l’ordre naturel des animaux composant les classes des Crustacés, des Arachnides et des Insectes; avec un tableau méthodique de leurs genre disposés en famille. Schoell, Paris, 444 pp.
- Linnaeus, C. (1758) Systema natura per regna tria naturae secundum classes, ordines, genera, species, cum characteribus, differentiis, synonymis, locis. Tenth edition. Laurentius Salvius, Holmiae, 824 pp.
- Liu, X.Y., Lu, X.M. & Zhang, W.W. (2016) New genera and species of the minute snakeflies (Raphidioptera: Mesoraphidiidae: Nanoraphidiini) from the mid Cretaceous of Myanmar. Zootaxa, 4103 (4), 301–324. http://doi.org/10.11646/zootaxa.4103.4.1
- Machado, R.J.P., Gillung, J.P., Winterton, S.L., Garzón‐Orduña, I.J., Lemmon, A.R., Lemmon, E.M. & Oswald, J.D. (2018) Owlflies are derived antlions: anchored phylogenomics supports a new phylogeny and classification of Myrmeleontidae (Neuroptera). Systematic Entomology, 44 (2), 418–450. https://doi.org/10.1111/syen.12334
- Makarkin, V.N. (2017) Oldest new genus of Myrmeleontidae (Neuroptera) from the Eocene Green River Formation, Colorado. Zootaxa, 4337 (4), 540–552. https://doi.org/10.11646/zootaxa.4337.4.5
- Makarkin, V.N. (2023) A new species of Mesoraphidiidae (Raphidioptera) from mid-Cretaceous Kachin amber, with discussion on anal veins in Raphidiomorpha. Cretaceous Research, 146, 105484. https://doi.org/10.1016/j.cretres.2023.105484
- 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. https://doi.org/10.1666/12-052R.1
- Makarkin, V.N. & Archibald, S.B. (2014) A revision of the late Eocene snakeflies (Raphidioptera) of the Florissant Formation, Colorado, with special reference to the wing venation of the Raphidiomorpha. Zootaxa, 3784 (4), 401–444. https://doi.org/10.11646/zootaxa.3784.4.4
- Makarkin, V.N. & Perkovsky, E.E. (2020) A new species of Proneuronema (Neuroptera, Hemerobiidae) from the late Eocene Rovno amber. Zootaxa, 4718 (2), 292–300. https://doi.org/10.11646/zootaxa.4718.2.11
- 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. http://doi.org/10.11646/zootaxa.4179.3.2
- Makarkin V.N., Archibald, S.B. & Jepson, J.E. (2019) The oldest Inocelliidae (Raphidioptera) from the Eocene of western North America. Canadian Entomologist, 151 (4), 521–530. https://doi.org/10.4039/tce.2019.26
- Makarkin, V.N., Antell, G. & Archibald, S.B. (2022) A revision of Chrysopidae (Neuroptera) of the late Eocene Florissant Formation, Colorado, with description of new species. Zootaxa, 5133 (3), 301–345. https://doi.org/10.11646/zootaxa.5133.3.1
- Martins, C.C., Flint, O.S. Jr. & Liu, X.Y. (2018) Notes on the pleasing lacewing genus Dilar Rambur, 1838 (Neuroptera, Dilaridae), with description of a new species from Vietnam. Zootaxa, 4425 (1), 193–200. http://doi.org/10.11646/zootaxa.4425.1.13
- McKenna, M. C. (1990) Plagiomenids (Mammalia: ?Dermoptera) from the Oligocene of Oregon, Montana and South Dakota, and Middle Eocene of northwestern Wyoming. In: Brown, T.M. & Rose, K.D. (Eds.), Dawn of the Age of Mammals in the Northern Part of the Rocky Mountain Interior, North America. Geological Society of America, Special Paper 243. Geological Society of America, Boulder, Colorado, pp. 211–234.
- Navás, L. (1909) Monografía de la familia de los Diláridos (Ins. Neur.). Memorias de la Real Academia de Ciencias y Artes de Barcelona, Series 3, 7, 619–671.
- Navás, L. (1910) Crisópidos (Ins. Neur.) nuevos. Brotéria, Zoológica, 9, 38–59.
- Navás, L. (1913) Neurópteros del R. Museo Zoológico de Nápoles. Annuario del [R.] Museo Zoologico della R. Università di Napoli, New Series, 4 (3), 1–11.
- Navás, L. (1914) Neurópteros de la Tripolitania. II serie. Annali del Museo Civico di Storia Naturale Giacomo Doria, 46, 202–209.
- Navás, L. (1916) Notas sobre el orden de los Rafidiópteros (Ins.). Memorias de la Real Academia de Ciencias y Artes de Barcelona, Series 3, 12, 507–513.
- Newman, E. (1838) Entomological Notes. Entomological Magazine, 5, 168–181 + 372–402, 483–500.
- Oswald, J.D. (2021) Neuropterida Species of the World. Lacewing Digital Library, Research Publication No. 1. Available from: http://lacewing.tamu.edu/SpeciesCatalog/Main (accessed 18 January 2023)
- Penny, N.D. (1982) Review of the generic level classification of the New World Ascalaphidae (Neuroptera). Acta Amazonica, 11, 391–406. [for 1981]
- Penny, N.D. (2002) Family Ascalaphidae. In: Penny, N.D. (Ed.), A Guide to the Lacewings (Neuroptera) of Costa Rica. Proceedings of the California Academy of Sciences, Series 4, 53, 176–186 (text), 293–299 (figures).
- Perkovsky, E.E. & Makarkin, V.N. (2019) A new species of Succinoraphidia Aspöck & Aspöck, 2004 (Raphidioptera: Raphidiidae) from the late Eocene Rovno amber, with venation characteristics of the genus. Zootaxa, 4576 (3), 570–580. https://doi.org/10.11646/zootaxa.4576.3.9
- Pierce, H.G. & Constenius, K.N. (2014) Terrestrial and aquatic mollusks of the Eocene Kishenehn Formation, Middle Fork Flathead River, Montana. Annals of the Carnegie Museum, 82, 305–329. https://doi.org/10.2992/007.082.0401
- Rambur, J.P. (1838) s.n. In: Faune entomologique de l’Andalousie. Vol. 2. A. Bertrand, Paris, pl. 9.
- Rambur, J.P. (1842) Histoire Naturelle des Insectes, Névroptères. Librairie encyclopédique de Roret. Fain et Thunot, Paris, [xviii] + 534 pp.
- Riek, E.F. (1968) A new genus and key to the species of Australian Stilbopterygidae (Neuroptera). Journal of the Australian Entomological Society, 7, 105–108.
- Schneider, W.G. (1851) Symbolae ad monographiam generis Chrysopae, Leach. Hirt, Vratislaviae, 178 pp.
- 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. https://doi.org/10.3133/70159191
- 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. https://doi.org/10.5962/bhl.title.44698
- Smith, J.B. (1900) Insects of New Jersey: a list of species occurring in New Jersey, with notes on those of economic importance. Annual Report of the New Jersey State Board of Agriculture, 27 (Supplement), 1–755. [for 1899]
- 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.
- van der Weele, H.W. (1903) Description of a new genus and species of holophthalmous Ascalaphidae. Notes from the Leyden Museum, 23, 234–236.
- van der Weele, H.W. (1909) Ascalaphiden. Collections Zoologiques du Baron Edm. de Selys Longchamps. Catalogue Systématique et Descriptif, 8 (for 1908), 1–326.
- Wilf, P., Wing, S.L., Greenwood, D.R. & Greenwood, C.L. (1998) Using fossil leaves as paleoprecipitation indicators: An Eocene example, Geology, 26, 203–206. https://doi.org/10.1130/0091-7613(1998)026%3C0203:UFLAPI%3E2.3.CO;2
- Winterton, S.L., Lemmon, A.R., Gillung, J.P., Garzon, I.J., Badano, D., Bakkes, D.K., Breitkreuz, L.C.V., Engel, M.S., Lemmon, E.M., Liu, X., Machado, R.J.P., Skevington, J.H. & Oswald, J.D. (2018). Evolution of lacewings and allied orders using anchored phylogenomics (Neuroptera, Megaloptera, Raphidioptera). Systematic Entomology, 43, 330–354. https://doi.org/10.1111/syen.12278