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Type: Article
Published: 2021-04-01
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Early Eocene snakeflies (Raphidioptera) of western North America from the Okanagan Highlands and Green River Formation

Department of Biological Sciences, Simon Fraser University, 8888 University Drive, Burnaby, British Columbia, V5A 1S6, Canada.
Federal Scientific Centre of the East Asia Terrestrial Biodiversity, Far Eastern Branch of the Russian Academy of Sciences, Vladivostok, Russia
Raphidioptera Raphidiidae early Eocene western North America

Abstract

Four new species of Raphidiidae are described from the early Eocene of western North America: Megaraphidia antiquissima sp. nov. from McAbee, M. ootsa sp. nov. from Driftwood Canyon, M. hopkinsi sp. nov. from the Allenby Formation (all from British Columbia, Canada), M. klondika sp. nov. from Republic (Washington, United States of America). Archiinocellia Handlirsch, 1910, Archiinocellia oligoneura Handlirsch, 1910 from Horsefly River (British Columbia, Canada), and A. protomaculata (Engel, 2011), comb. nov., from the Green River Formation (Colorado, United States of America) are redescribed. Archiinocellia is assigned to Raphidiidae, sit. nov. The apparent absence of sclerotized gonocoxites 9 in the Archiinocellia protomaculata male is probably plesiomorphic at the family level. As some modern snakeflies do not require a cold interval to complete their development and Eocene Inocelliidae and Raphidiidae lived in regions of warm winters (especially A. protomaculata), adaptation to cold winters in many modern snakeflies is a post-Eocene phenomenon. Eocene Raphidiidae of Europe (Priabonian) differ greatly from those of North America (Ypresian and Priabonian). This pattern might reflect dispersal in either direction or ranges established prior to continental separation. Eocene Inocelliidae of Europe (Priabonian), however, are more similar to those of North America (Ypresian and Lutetian).

 

References

  1. Abbt, V., Gruppe, A., Aspöck, H. & Aspöck, U. (2018) Einfluss der Überwinterungsdauer auf die Metamorphose bei Raphidioptera-Larven. Mitteilungen der Deutschen Gesellschaft für Allgemeine und Angewandte Entomologie, 21, 305–308.

    AEMET (2020) La Agencia Estatal de Meteorología. Online. [http://www.aemet.es/en/serviciosclimaticos/datosclimatologicos]

    Albarda, H. (1891) Révision des Rhaphidides. Tijdschrift voor Entomologie, 34, 65–184.

    Allen, S.E., Lowe, A.J., Peppe, D.J. & Meyer, H.W. (2020) Paleoclimate and paleoecology of the latest Eocene Florissant flora (Central Colorado, USA). Palaeogeography, Palaeoclimatology, Palaeoecology, 551, 109678.

    https://doi.org/10.1016/j.palaeo.2020.109678 .

    Antell, G.S. & Kathirithamby, J. (2016) The first twisted-wing parasitoids (Insecta: Strepsiptera) from the Early Eocene Green River Formation of Colorado. Bulletin of the Peabody Museum of Natural History, 57, 165–174.

    https://doi.org/10.3374/014.057.0204

    Archibald, S.B. (2005) New Dinopanorpidae (Insecta: Mecoptera) from the Eocene Okanagan Highlands (British Columbia, Canada; Washington State, USA). Canadian Journal of Earth Sciences, 42, 119–136.

    https://doi.org/10.1139/e04-073

    Archibald, S.B., Bossert, W.H., Greenwood, D.R. & Farrell, B.D. (2010) Seasonality, the latitudinal gradient of diversity, and Eocene insects. Paleobiology, 36, 374–398.

    https://doi.org/10.1666/09021.1

    Archibald, S.B. & Farrell, B.D. (2003) Wheeler’s Dilemma. Proceedings of the Second Paleoentomological Congress. Acta Zoologica Crakoviensia, 46(suppl.), 17–23.

    Archibald, S.B., Greenwood, D.R. & Mathewes, R.W. (2013a) Seasonality, montane beta diversity, and Eocene insects: testing Janzen’s dispersal hypothesis in an equable world. Palaeogeography, Palaeoclimatology, Palaeoecology, 371, 1–8.

    https://doi.org/10.1016/j.palaeo.2012.10.043

    Archibald, S.B., Greenwood, D.R., Smith, R.Y., Mathewes, R.W. & Basinger, J.F. (2011) Early Eocene lagerstätten of the Okanagan Highlands (British Columbia and Washington State). Geoscience Canada, 38, 145–154.

    Archibald, S.B., Mathewes, R.W. & Greenwood, D.R. (2013b) The Eocene apex of panorpoid scorpionfly family diversity. Journal of Paleontology, 87, 677–695.

    https://doi.org/10.1666/12-129

    Archibald, S.B., Morse, G.E., Greenwood, D.R. & Mathewes, R.W. (2014) Fossil palm beetles refine upland winter temperatures in the Early Eocene Climatic Optimum. Proceedings of the National Academy of Sciences of the United States of America, 111, 8095–8100.

    https://doi.org/10.1073/pnas.1323269111

    Archibald, S.B. & Makarkin, V.N. (2020) A new genus and species of split-footed lacewings (Neuroptera) from the early Eocene of western Canada and revision of the subfamily affinities of Mesozoic Nymphidae. The Canadian Entomologist, 152, 269–287.

    https://doi.org/10.4039/tce.2020.10

    Archibald, S.B., Makarkin, V.N. & Ansorge, J. (2009) New fossil species of Nymphidae (Neuroptera) from the Eocene of North America and Europe. Zootaxa, 2157 (1), 59–68.

    https://doi.org/10.11646/zootaxa.2157.1.4

    Archibald, S.B., Rasnitsyn, A.P., Brothers, D.J. & Mathewes, R.W. (2018) Modernisation of the Hymenoptera: ants, bees, wasps and sawflies of the early Eocene Okanagan Highlands. The Canadian Entomologist, 150, 205–257.

    https://doi.org/10.4039/tce.2017.59

    Aspöck, H. (1998) Distribution and biogeography of the order Raphidioptera: updated facts and a new hypothesis. Acta Zoologica Fennica, 209, 33–44.

    Aspöck, H. (2000) Der endkreidezeitliche Impakt und das Überleben der Raphidiopteren. Entomologica Basiliena, 22, 223–233.

    Aspöck, H. (2002) The biology of Raphidioptera: a review of present knowledge. In: G. Sziráki, G. (Ed.), Neuropterology 2000. Proceedings of the Seventh International Symposium on Neuropterology. Acta Zoologica Academiae Scientiarum Hungaricae, 48 (Supplement 2), 35–50.

    Aspöck, H. (2004) Kamelhalsfliegen—lebende Fossilien: eine der Endkreide-Katastrophe entkommene Tiergruppe. In: Verhandlungen Westdeutscher Entomologentag, Düsseldorf, 2002, pp. 1–6.

    Aspöck, H., Aspöck, U. & Rausch, H. (1991) Die Raphidiopteren der Erde. Eine monographische Darstellung der Systematik, Taxonomie, Biologie, Ökologie und Chorologie der rezenten Raphidiopteren der Erde, mit einer zusammenfassenden Übersicht der fossilen Raphidiopteren (Insecta: Neuropteroidea). 2 Vols. Goecke & Evers, Krefeld, 730 and 550 pp.

    Aspöck, U. (1975) The present state of knowledge on the Raphidioptera of America (Insecta: Neuropteroidea). Polskie Pismo Entomologiczne, 45, 537–546.

    Aspöck, U. (1982) Polymorphismus und Polytypie bei nearktischen Raphidiiden: Agulla (Glavia) modesta Carpenter, Agulla (Glavia) unicolor Carpenter und Agulla (Glavia) paramerica n. sp. (Neuropteroidea: Raphidioptera). Zeitschrift der Arbeitsgemeinschaft Österreichischer Entomologen, 33, 95–5112. [for 1981]

    Aspöck, U. (1987) Agulla faulkneri n. sp.—eine neue Raphidiiden-Spezies aus Kalifornien (Neuropteroidea: Raphidioptera: Raphidiidae). Zeitschrift der Arbeitsgemeinschaft Österreichischer Entomologen, 39, 1–4.

    Aspöck, U. (1988) Negha meridionalis n. sp. – eine neue Inocelliiden-Spezies aus Süd-Kalifornien (U.S.A.) und Baja California (Mexiko) (Neuropteroidea: Raphidioptera: Inocelliidae). Zeitschrift der Arbeitsgemeinschaft Österreichischer Entomologen, 39, 107–112. [for 1987]

    Aspöck, U. & Aspöck, H. (1970) Untersuchungen über die Raphidiopteren Mexikos (Insecta, Neuropteroidea). Polskie Pismo Entomologiczne, 40, 705–725.

    Aspöck, U. & Aspöck, H. (1978) Raphidia (Aliaberaphidia) tenochtitlana n. sp.—eine neue Raphidiiden-Spezies aus Baja California (Mexiko) (Neuropteroidea, Raphidioptera). Zeitschrift der Arbeitsgemeinschaft Österreichischer Entomologen, 30, 21–24.

    Aspöck, U. & Aspöck, H. (1994) Paradoxe Verbreitungsbilder von Neuropteroidea (Insecta: Raphidioptera, Neuroptera). Zeitschrift der Arbeitsgemeinschaft Österreichischer Entomologen, 46, 30–44.

    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, U. & Aspöck, H. (2013) Alena Navás, 1916 – the dethroned genus and Alena (Aztekoraphidia) michoacana sp. n. from Mexico (Neuropterida: Raphidioptera: Raphidiidae). Deutsche Entomologische Zeitschrift, 60, 53–58.

    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., Aspöck, H. & Rausch, H. (1994a) Alena (Mexicoraphidia) americana (Carpenter, 1958): Taxonomie, Systematik, Ökologie und Chorologie (Neuropteroidea: Raphidioptera: Raphidiidae). Zeitschrift der Arbeitsgemeinschaft Österreichischer Entomologen, 46, 131–139.

    Aspöck, U., Aspöck, H. & Rausch, H. (1994b) Neue Arten der Familie Raphidiidae aus Mexiko und Nachweis einer Spermatophore in der Ordnung Raphidioptera (Insecta: Neuropteroidea). Entomologia Generalis, 18, 145–163.

    https://doi.org/10.1127/entom.gen/18/1994/145

    Aspöck, U., Haring, E. & Aspöck, H. (2012a) Biogeographical implications of a molecular phylogeny of the Raphidiidae (Raphidioptera). Mitteilungen der Deutschen Gesellschaft fur allgeimeine und angewandte Entomologie, 18, 575–582.

    Aspöck, U. & Contreras-Ramos, A. (2004) Alena (Aztekoraphidia) horstaspoecki nov. spec. - a new snakefly from Mexico (Raphidioptera, Raphidiidae). Denisia, 13, 129–134.

    Aspöck, H., Liu, X.Y. & Aspöck, U. (2012b) The family Inocelliidae (Neuropterida: Raphidioptera): A review of present knowledge. Mitteilungen der Deutschen Gesellschaft für Allgemeine und Angewandte Entomologie, 18, 565–573.

    Aspöck, U., Liu, X.Y., Rausch, H. & Aspöck, H. (2011) The Inocelliidae of southeast Asia: a review of present knowledge (Raphidioptera). Deutsche Entomologische Zeitschrift, (N.F.), 58, 259–274.

    https://doi.org/10.1002/mmnd.201100029

    Banks, N. (1911) Descriptions of new species of North American neuropteroid insects. Transactions of the American Entomological Society, 37, 335–360.

    Barton, D.G. & Wilson, M.V.H. (2005) Taphonomic variations in Eocene fish-bearing varves at Horsefly, British Columbia, reveal 10 000 years of environmental change. Canadian Journal of Earth Sciences, 42, 137–149.

    https://doi.org/10.1139/e05-001

    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.

    https://doi.org/10.1016/S0016-7878(09)80005-9

    Blades, D.C.A. (2019) Raphidioptera of Canada. ZooKeys, 819, 383–386.

    https://doi.org/10.3897/zookeys.819.26626

    Brikiatis, L. (2014) The De Geer, Thulean and Beringia routes: key concepts for understanding early Cenozoic biogeography. Journal of Biogeography, 41, 1036–1054.

    https://doi.org/10.1111/jbi.12310

    Carpenter, F.M. (1936) Revision of the Nearctic Raphidiodea (recent and fossil). Proceedings of the American Academy of Arts and Science, 71, 89–157.

    https://doi.org/10.2307/20023217

    Carpenter, F.M. (1957) The Baltic amber snake-flies Neuroptera. Psyche, 63(for 1956), 77–81.

    https://doi.org/10.1155/1956/97248

    Carpenter, F.M. (1958) Mexican snake-flies (Neuroptera: Raphidiodea). Psyche, 65, 52–58.

    https://doi.org/10.1155/1958/14859

    Carpenter, F.M. (1992) Superclass Hexapoda. In: Kaesler, R.L. (Ed.), Treatise on Invertebrate Paleontology. Part R. Arthropoda 4. Vols. 3 & 4. The Geological Society of America, Boulder, Colorado & University of Kansas, Lawrence, Kansas, pp. 1–655.

    Cockerell, T.D.A. (1907) Some fossil arthropods from Florissant, Colorado. Bulletin of the American Museum of Natural History, 23, 605–616.

    Cockerell, T.D.A. (1909a) Fossil insects from Florissant, Colorado. Bulletin of the American Museum of Natural History, 26, 67–76.

    Cockerell, T.D.A. (1909b) 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. (1912) A fossil Raphidia (Neur. Planip.). Entomological News, 23, 215–216.

    Cockerell, T.D.A. (1914) New and little-known insects from the Miocene of Florissant, Colorado. The Journal of Geology, 22, 714–724.

    https://doi.org/10.1086/622186

    Cockerell, T.D.A. (1917) Some fossil insects from Florissant, Colorado. Proceedings of the United States National Museum, 53 (2210), 389–392.

    https://doi.org/10.5479/si.00963801.2210.389

    Cockerell, T.D.A. & Custer, C. (1925) A new fossil Inocellia from Florissant. The Entomologist, 58, 295–297.

    Codington, L.A. (1995) Note on a possible paleospiderweb from Douglas Pass, Colorado. In: Averett, W.R. (Ed.), The Green River Formation in Piceance Creek and Eastern Uinta Basins. Grand Junction Geological Society, Grand Junction, Colorado, pp. 117–118.

    Condamine, F.L., Clapham M.E. & Kergoat G.J. (2016) Global patterns of insect diversification: towards a reconciliation of fossil and molecular evidence? Scientific Reports, 6, 19208.

    https://doi.org/10.1038/srep19208

    Constenius, K.N. (1996) Late Paleogene extensional collapse of the Cordilleran foreland fold and thrust belt. Geological Society of America Bulletin, 108, 20–39.

    https://doi.org/10.1130/0016-7606(1996)108<0020:LPECOT>2.3.CO;2

    Dayvault, R.D., Codington, L.A., Kohls, D., Hawes, W.D. & Ott, P.M. (1995) Fossil insects and spiders from three locations in the Green River Formation of the Piceance Creek Basin, Colorado. In: Averett, W.R. (Ed.), The Green River Formation in Piceance Creek and Eastern Uinta Basins. Grand Junction Geological Society, Grand Junction, Colorado, pp. 97–115.

    Engel, M.S. (1998) A new fossil snake-fly species from Baltic amber (Raphidioptera: Inocellidae). Psyche, 102, 187–193. [for 1995]

    https://doi.org/10.1155/1995/23626

    Engel, M.S. (2002) The smallest snakefly (Raphidioptera: Mesoraphidiidae): A new species in Cretaceous amber from Myanmar, with a catalog of fossil snakeflies. American Museum Novitates, 3363, 1–22.

    Engel, M.S. (2003) A new Eocene-Oligocene snakefly from Florissant, Colorado (Raphidioptera: Raphidiidae). Transactions of the Kansas Academy of Sciences, 106, 124–128.

    Engel, M.S. (2009) A Miocene snakefly from Stewart Valley, Nevada (Raphidioptera: Raphidiidae). Transactions of the Kansas Academy of Science, 112, 211–214.

    https://doi.org/10.1660/062.112.0409

    Engel, M.S. (2011) A new snakefly from the Eocene Green River Formation (Raphidioptera: Raphidiidae). Transactions of the Kansas Academy of Science, 114, 77–87.

    https://doi.org/10.1660/062.114.0107

    Ewing, T. (1980) Paleogene tectonic evolution of the Pacific Northwest. The Journal of Geology, 88, 619–638.

    https://doi.org/10.1086/628551

    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. & Johnson, K. (Eds.), Fossil flora and stratigraphy of the Florissant Formation, Colorado. Proceedings of the Denver Museum of Nature and Science, Series 4, 1, pp. 1–16.

    Greenwood, D.R., Archibald, S.B., Mathewes, R.W. & Moss, P.T. (2005) Fossil biotas from the Okanagan Highlands, southern British Columbia and northeastern Washington State: climates and ecosystems across an Eocene landscape. Canadian Journal of Earth Sciences, 42, 167–185.

    https://doi.org/10.1139/e04-100

    Grimaldi, D. & Engel, M.S. (2005) Evolution of the insects. Cambridge University Press, New York, xv + 755 pp.

    Gröhn, C. (2015) Einschlüsse im baltischen Bernstein. Wachholtz Verlag—Murmann, Kiel, 424 pp.

    Handlirsch, A. (1910) Contributions to Canadian Palaeontology. Vol. II. Part III. Canadian fossil insects. V. Insects from the Tertiary lake deposits of the Southern Interior of British Columbia collected by Mr. Lawrence Lambe, in 1906. Memoirs of the Geological Survey of Canada, 12, 93–129.

    https://doi.org/10.4095/100486

    Haring, E., Aspöck, H., Bartel, D. & Aspöck, U. (2011) Molecular phylogeny of the Raphidiidae (Raphidioptera). Systematic Entomology, 36, 16–30.

    https://doi.org/10.1111/j.1365-3113.2010.00542.x

    Janzen, J.-W. (2002) Arthropods in Baltic amber. Ampyx Verlag, Halle, 167 pp.

    Jarzembowski, E.A. (1989) Cretaceous insect extinction. Mesozoic Research, 2, 25–38.

    Kovarik, P.W., Burke, H.R. & Agnew, C.W. (1991) Development and behavior of a snakefly, Raphidia bicolor Albarda (Neuroptera: Raphidiidae). Southwestern Entomologist, 16, 353–364.

    Labandeira, C.C., Johnson, K.R. & Wilf, P. (2002) Impact of the terminal Cretaceous event on plant—insect associations. Proceedings of the National Academy of Sciences of the United States of America, 99, 2061–2066.

    https://doi.org/10.1073/pnas.042492999

    Labandeira, C.C. & Sepkoski, J.J., Jr. (1993) Insect diversity in the fossil record. Science, 261, 310–315.

    https://doi.org/10.1126/science.11536548

    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.

    Liu, X.Y., Aspöck, H., Yang, D. & Aspöck, U. (2010) Species of the Inocellia fulvostigmata group (Raphidioptera, Inocelliidae) from China. Deutsche Entomologische Zeitschrift (N.F.), 57, 223–232.

    https://doi.org/10.1002/mmnd.201000019

    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, 301–324.

    https://doi.org/10.11646/zootaxa.4103.4.1

    Liu, X.Y., Makarkin, V.N., Yang, Q. & Ren, D. (2013) A remarkable new genus of basal snakeflies (Insecta: Raphidioptera: Priscaenigmatomorpha) from the Early Cretaceous of China. Cretaceous Research, 45, 306–313.

    https://doi.org/10.1016/j.cretres.2013.06.001

    Liu, X.Y., Ren, D. & Yang, D. (2014) New transitional fossil snakeflies from China illuminate the early evolution of Raphidioptera. BMC Evolutionary Biology, 14, 84.

    https://doi.org/10.1186/1471-2148-14-84

    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., Archibald, S.B. & Jepson, J.E. (2019) The oldest Inocelliidae (Raphidioptera) from the Eocene of western North America. The Canadian Entomologist, 151, 521–530.

    https://doi.org/10.4039/tce.2019.26

    Manchester, S.R. (2001) Update on the Megafossil Flora of Florissant, Colorado. In: Evanoff, E., Gregory-Wodzicki, K. & Johnson, K. (Eds.), Fossil flora and stratigraphy of the Florissant Formation, Colorado. Proceedings of the Denver Museum of Nature and Science, Series 4, 1, 137–162.

    Martynov, A.V. (1925) To the knowledge of fossil insects from Jurassic beds in Turkestan. 1. Raphidioptera. Bulletin of the Academy of Sciences USSR, Series 6, 19, 233–246.

    Monserrat, V.J. & Papenberg, D. (2006) Revisión del género Harraphidia Steinmann, 1963 con la descripción de dos nuevas especies de la península Ibérica y de Marruecos (Insecta, Raphidioptera). Graellsia, 62, 203–222.

    https://doi.org/10.3989/graellsia.2006.v62.i2.67

    Moss, P.T., Greenwood, D.R. & Archibald, S.B. (2005) Regional and local vegetation community dynamics of the Eocene Okanagan Highlands (British Columbia-Washington State) from palynology. Canadian Journal of Earth Sciences, 42, 187–204.

    https://doi.org/10.1139/e04-095

    Navás, L. (1909). Rhaphidides (Insectes Névroptères) du Musée de Paris. Annales de la Société Scientifique de Bruxelles, 33 (1), 143–146.

    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. (1914a) Algunos Neurópteros del Museo de Oxford. I serie. Boletín de la Sociedad Aragonesa de Ciencias Naturales, 13, 61–68.

    Navás, L. (1914b) Névroptères nouveaux de l’Amerique du Nord. IIème série. Entomologische Zeitschrift (Frankfurt am Main), 28,18–20 + 25–26.

    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.

    Nel, A. (1993) Nouveaux Raphidioptères fossiles du Cénozoïque de France et d’Espagne (Raphidioptera, Raphidiidae, Inocelliidae). Ecole Pratique des Hautes Etudes, Biologie et Evolution des Insectes, 6, 99–108.

    Oswald, J.D. (1990) Raphidioptera. In: Grimaldi, D.A. (Ed.), Insects from the Santana Formation, Lower Cretaceous, of Brazil. Bulletin of the American Museum of Natural History, 195, 154–163.

    Oswald, J.D. (1993) Revision and cladistic analysis of the world genera of the family Hemerobiidae (Insecta: Neuroptera). Journal of the New York Entomological Society, 101, 143–299.

    Palmer, A.R. & Strobeck, C. (1986) Fluctuating asymmetry: measurement, analysis, patterns. Annual Review of Ecological Systematics, 17, 391–421.

    https://doi.org/10.1146/annurev.es.17.110186.002135

    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

    Perkovsky, E.E., Rasnitsyn, A.P., Vlaskin, A.P. & Taraschuk, M.V. (2007) A comparative analysis of the Baltic and Rovno amber arthropod faunas: representative samples. African Invertebrates, 48, 229–245.

    Pictet-Baraban, F.J. & Hagen, H.A. (1856) Die im Bernstein befindlichen Neuropteren der Vorwelt. In: G.C. Berendt (Ed.), Die im Bernstein befindlichen organischen Reste der Vorwelt gesammelt, in verbindung mit mehreren bearbeitet und herausgegeben von Dr. Georg Carl Berendt. Band 2. Abteilung 2. Nicholaischen Buchhandlung, Berlin, pp. 41–125.

    Pierce, H. & Constenius, K.N. (2001) Late Eocene-Oligocene nonmarine mollusks of the northern Kishenehn Basin, Montana and British Columbia. Annals of the Carnegie Museum, 70, 1–112.

    Pribyl, L.J., Labandeira, C.C. & Kohls, S.D. (1998) Biodiversity and taphonomy of fossil insects from the Green River Formation, Piceance Creek Basin, Colorado, USA. In: First International Paleoentomological Conference, Moscow, Russia, 30 August–4 September 1998, Abstracts, pp. 37.

    Rasnitsyn, A.P. (2002) Introduction to palaeoentomology. In: Rasnitsyn, A.P. & Quicke, D.L.J. (Eds.), History of Insects. Kluwer Academic Publishers, Dordrecht, pp. 1–63.

    https://doi.org/10.1007/0-306-47577-4

    Rasnitsyn, A.P., Bashkuev, A.S., Kopylov, D.S., Lukashevich, E.D., Ponomarenko, A.G., Popov, Y.A., Rasnitsyn, D.A., Ryzhkova, O.V., Sidorchuk, E.A., Sukatsheva, I.D. & Vorontsov, D.D. (2016) Sequence and scale of changes in the terrestrial biota during the Cretaceous (based on materials from fossil resins). Cretaceous Research, 61, 234–255.

    https://doi.org/10.1016/j.cretres.2015.12.025

    Read, P. (2000) Geology and Industrial Minerals of the Tertiary Basins, South-Central British Columbia. British Columbia Ministry of Energy and Mines, GeoFile 2000-3, 1–102.

    Rohwer, S.A. (1909) Three new fossil insects from Florissant, Colorado. American Journal of Science, 28, 533–536.

    https://doi.org/10.2475/ajs.s4-28.168.533

    Rouse, G.E., Hopkins, W.S., Jr. & Piel, K.M. (1970) Palynology of some Late Cretaceous and Early Tertiary deposits in British Columbia and adjacent Alberta. Geological Society of America, Special Paper, 127, 213–246.

    https://doi.org/10.1130/SPE127-p213

    Rust, J. (1999) Biologie der Insekten aus dem ältesten Tertiär Nordeuropas. Habilitationsschrift zur Erlangung der venia legendi für das Fach Zoologie in der biologischen Fakultät der Georg-August-Universitäat Göttingen, Göttingen, 482 pp.

    Scudder, S.H. (1890) The Tertiary Insects of North America. Report of the United States Geological Survey of the Territories, 13, 1–734.

    https://doi.org/10.5962/bhl.title.44698

    Smith, M.E., Carroll, A. & Singer, B. (2008) Synoptic reconstruction of a major ancient lake system: Eocene Green River Formation, western United States. Geological Society of America Bulletin, 120, 54–84.

    https://doi.org/10.1130/B26073.1

    Smith, M.E., Singer, B. & Carroll, A. (2003) 40Ar/39Ar geochronology of the Green River Formation, Wyoming. Bulletin of the Geological Society of America, 115, 549–565.

    https://doi.org/10.1130/0016-7606(2003)115<0549:AGOTEG>2.0.CO;2

    Tribe, S. (2005) Eocene paleo-physiography and drainage directions, southern Interior Plateau, British Columbia. Canadian Journal of Earth Sciences, 42, 215–230.

    https://doi.org/10.1139/e04-062

    Wang, W.L. (1987) Early Mesozoic insect fossils from western Liaoning. In: Yu, X.H., Wang, W.L., Liu, X.T., Zhang, W., Zheng, S.L., Yu, J.H., Ma, F.Z., Dong, G.Y. & Yao, P.Y. (Eds.), Mesozoic stratigraphy and palaeontology of western Liaoning. Vol. 3. Geological Publishing House, Beijing, pp. 202–222. [in Chinese, English summary]

    Weitschat, W. & Wichard, W. (1998) Atlas der Pflanzen und Tiere im Baltischen Bernstein. Dr. Friedrich Pfeil Verlag, München, 256 pp.

    Wehr, W.C. & Barksdale, L.L. (1996) A checklist of fossil insects from Republic, Washington. Washington Geology, 24, 29.

    Whalley, P.E.S. (1985) The systematics and palaeogeography of the Lower Jurassic insects of Dorset, England. Bulletin of the British Museum of Natural History, Geology, 39, 107–189.

    Wichard, W., Chatterton, C. & Ross A. (2005) Corydasialidae fam. n. (Megaloptera) from Baltic amber. Insect Systematics and Evolution, 36, 279–283.

    https://doi.org/10.1163/187631205788838410

    Wilf, P., Labandeira, C.C., Johnson, K.R. & Ellis, B. (2006) Decoupled plant and insect diversity after the end-Cretaceous extinction. Science, 313, 1112–1115.

    https://doi.org/10.1126/science.1129569

    Willmann, R. (1994) Raphidiodea aus dem Lias und die Phylogenie der Kamelhalsfliegen (Insecta: Holometabola). Paläontologische Zeitschrift, 68, 167–197.

    https://doi.org/10.1007/BF02989439

    Wilson, M.V.H. (1977a) Paleoecology of Eocene lacustrine varves at Horsefly, British Columbia. Canadian Journal of Earth Sciences, 14, 953–962.

    https://doi.org/10.1139/e77-089

    Wilson, M.V.H. (1977b) New records of insect families from the freshwater Middle Eocene of British Columbia. Canadian Journal of Earth Sciences, 14, 1139–1155.

    https://doi.org/10.1139/e77-104

    Woglum, R.S. & McGregor, E.A. (1959) Observations on the life history and morphology of Agulla astuta (Banks) (Neuroptera: Raphidiodea: Raphidiidae). Annals of the Entomological Society of America, 52, 489–502.

    https://doi.org/10.1093/aesa/52.5.489

    Wolfe, J.A., Gregory-Wodzicki, K.M., Molnar, P. & Mustoe, G. (2003) Rapid uplift and then collapse in the Eocene of the Okanagan? Evidence from paleobotany. In: Geological Association of Canada—Mineralogical Association of Canada—Society of Economic Geologists, Joint Annual Meeting, Vancouver, Abstracts, 28, 533. [CD-ROM]

    Young, R.G. (1995) Stratigraphy of Green River Formation in Piceance Creek Basin, Colorado. In: Averett, W.R. (Ed.), The Green River Formation in Piceance Creek and Eastern Uinta Basins. Grand Junction Geological Society, Grand Junction, Colorado, pp. 1–13.

    Zachos, J.C., Dickens, G.R. & Zeebe, R.E. (2008) An early Cenozoic perspective on greenhouse warming and carbon-cycle dynamics. Nature, 451, 279–283.

    https://doi.org/10.1038/nature06588

    Zeleny, J. (1969) Variability in the species Raphidia ophiopsis Linnaeus, and Agulla xanthostigma (Schummel) (Raphidioptera). Acta Entomologica Bohemoslovaca, 66, 15–38.

    Zherikhin, V.V. (1978) Development and changes in Cretaceous and Cenozoic faunistic complexes (tracheates and chelicerates). Trudy Paleontologicheskogo Instituta, 165, pp. 1–200. [in Russian]