Abstract
We describe the Cephalozygoptera, a new, extinct suborder of Odonata, composed of the families Dysagrionidae and Sieblosiidae, previously assigned to the Zygoptera, and possibly the Whetwhetaksidae n. fam. The Cephalozygoptera is close to the Zygoptera, but differs most notably by distinctive head morphology. It includes 59 to 64 species in at least 19 genera and one genus-level parataxon. One species is known from the Early Cretaceous (Congqingia rhora Zhang), possibly three from the Paleocene, and the rest from the early Eocene through late Miocene. We describe new taxa from the Ypresian Okanagan Highlands of British Columbia, Canada and Washington, United States of America: 16 new species of Dysagrionidae of the existing genus Dysagrion (D. pruettae); the new genera Okanagrion (O. threadgillae, O. hobani, O. beardi, O. lochmum, O. angustum, O. dorrellae, O. liquetoalatum, O. worleyae, all new species); Okanopteryx (O. jeppesenorum, O. fraseri, O. macabeensis, all new species); Stenodiafanus (S. westersidei, new species); the new genus-level parataxon Dysagrionites (D. delinei new species, D. sp. A, D. sp. B, both new); and one new genus and species of the new family Whetwhetaksidae (Whetwhetaksa millerae).
References
Allena, 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. [in press]
https://doi.org/10.1016/j.palaeo.2020.109678
Archibald, S.B. (2007) Climate and Species Diversity: The Eocene Okanagan Highlands Insect View. Vol. 1. Ph.D. Thesis, Harvard University, Cambridge, Massachusetts, 362 pp.
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. & Cannings, R.A. (2019) Fossil dragonflies (Odonata: Anisoptera) from the early Eocene Okanagan Highlands, western North America. The Canadian Entomologist, 151, 783–816.
https://doi.org/10.4039/tce.2019.61
Archibald, S.B. & Farrell, B.D. (2003) Wheeler’s dilemma. Proceedings of the Second Paleoentomological Congress, Acta Zoologica Crakoviensia, 46 (Supplement), Fossil Insects, 17–23.
Archibald, S.B., Greenwood, D.R. & Mathewes, R.W. (2013) 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. (2011a) Early Eocene lagerstätten of the Okanagan Highlands (British Columbia and Washington State). Geoscience Canada, 38, 145–154.
Archibald, S.B., Johnson, K.R., Mathewes, R.W. & Greenwood, D.R. (2011b) Intercontinental dispersal of giant thermophilic ants across the Arctic during early Eocene hyperthermals. Proceedings of the Royal Society B, 278, 3679–3686.
https://doi.org/10.1098/rspb.2011.0729
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 Paleontology, 4, 119–155, 307 (errata).
https://doi.org/10.1017/S1477201906001817
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., 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
Bechly, G. (1996) Morphologische Untersuchungen am Flügelgeäder der rezenten Libellen und deren Stammgruppenvertreter (Insecta; Pterygota; Odonata) unter besonderer Berüchsichtigung der Phylogenetischen Systematik und des Grundplanes der Odonata. Petalura, Böblingen, Special Volume, 2, 1–402.
Bechly, G. (1998) New fossil damselflies from Baltic amber, with description of a new species, a redescription of Litheuphaea carpenteri Fraser, and a discussion on the phylogeny of Epallagidae (Zygoptera: Caloptera). International Journal of Odonatology, 1, 33–63.
https://doi.org/10.1080/13887890.1998.9748092
Bechly, G. (2015) Fossile Libellennachweise aus Deutschland (Odonatoptera). Libellula Supplement, 14, 423–464.
Bechly, G., Nel, A., Martínez-Delclòs, X. & Fleck, G. (1998) Four new dragonflies from the Upper Jurassic of Germany and the Lower Cretaceous of Mongolia (Anisoptera: Hemeroscopidae, Sonidae, and Proterogomphidae fam. nov.). Odonatologica, 7, 149–187.
Bechly, G., Nel, A., Martinez-Delclos, X., Jarzembowski, E.A., Coram, R., Martill, D., Fleck, G., Escuillie, F., Wisshak, M.M. & Maisch, M. (2001) A revision and phylogenetic study of Mesozoic Aeshnoptera, with description of numerous new taxa (Insecta: Odonata: Anisoptera). Neue Paläontologische Abhandlungen, 4, 1–219, 48 plates.
Bechly, G. & Poinar Jr., G. (2013) Burmaphlebia reifi gen. et sp. nov., the first anisozygopteran damsel-dragonfly (Odonata: Epiophlebioptera: Burmaphlebiidae fam. nov.) from Early Cretaceous Burmese amber. Historical Biology, 25, 233–237.
https://doi.org/10.1080/08912963.2012.753884
Blanke, A., Greve, C., Mokso, R., Beckmann, F. & Misof, B. (2013) An updated phylogeny of Anisoptera including formal convergence analysis of morphological characters. Systematic Entomology, 38, 474–490.
https://doi.org/10.1111/syen.12012
Breedlovestrout, R.L., Evraets, B.J. & Parrish, J.T. (2013) New Paleogene paleoclimate analysis of western Washington using physiognomic characteristics from fossil leaves. Palaeogeography, Palaeoclimatology, and Palaeoecology, 392, 22–40.
https://doi.org/10.1016/j.palaeo.2013.08.013
Büsse, S. (2016) Morphological re-examination of Epiophlebia laidlawi (Insecta: Odonata) including remarks on taxonomy. International Journal of Odonatology, 19, 221–238.
https://doi.org/10.1080/13887890.2016.1257442
Bybee, S.M., Ogden, T.H., Branham, M.A. & Whiting, M.F. (2008) Molecules, morphology and fossils: a comprehensive approach to odonate phylogeny and the evolution of the odonate wing. Cladistics, 24, 477-514.
https://doi.org/10.1111/j.1096-0031.2007.00191.x
Calvert, P.P. (1913) The fossil Odonate Phenacolestes, with a discussion of the venation of the legion Podagrion Selys. Proceedings of the Academy of Natural Sciences of Philadelphia, 65: 225–272.
Campion, H. (1913) The Antenodal Reticulation of the Wings of Agrionine Dragonflies. Proceedings of the Academy of Natural Sciences of Philadelphia, 65, 220–224.
Carle, F.L. (2012) A new Epiophlebia (Odonata: Epiophlebioidea) from China with a review of epiophlebian taxonomy, life history, and biogeography. Arthropod Systematics & Phylogeny, 70, 75–83.
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. (1908a) Fossil insects from Florissant, Colorado. Bulletin of the American Museum of Natural History, 24, 59–69, 1 pl.
Cockerell, T.D.A. (1908b) Descriptions of Tertiary Insects IV. American Journal of Science, 26, 69–75.
https://doi.org/10.2475/ajs.s4-26.151.69
Cockerell, T.D.A. (1908c) Some Results of the Florissant Expedition of 1908. The American Naturalist, 42, 569–581.
https://doi.org/10.1086/278976
Cockerell, T.D.A. (1927) A new fossil dragonfly from the Eocene of Colorado. The Entomologist, 60, 81–82.
De Bivort, B.L., Clouse, R.M. & Giribet, G. (2010) A morphometrics-based phylogeny of the temperate Gondwanan mite harvestmen (Opiliones, Cyphophthalmi, Pettalidae). Journal of Zoological Systematics and Evolutionary Research, 48, 294–309.
https://doi.org/10.1111/j.1439-0469.2009.00562.x
Dijkstra, K.-D.B., Bechly, G., Bybee, S.M., Dow, R.A., Dumont, H.J., Fleck, G., Garrison, R.W., Hämäläinen, M., Kalkman, V.J., Karube, H., May, M.L., Orr, A.G., Paulson, D.R., Rehn, A.C., Theischinger, G., Trueman, J.W.H., van Tol, J., von Ellenrieder, N. & Ware, J. (2013) The classification and diversity of dragonflies and damselflies (Odonata). In: Animal biodiversity: an outline of higher-level classification and survey of taxonomic richness. In: Zhang, Z.-Q. (Ed.), Zootaxa, 3703, pp. 36–45.
https://doi.org/10.11646/zootaxa.3703.1.9
Dijkstra, K.-D.B., Kalkman, V.J., Dow, R.A., Stokvis, F.R. & van Tol, J. (2014) Redefining the damselfly families: the first comprehensive molecular phylogeny of Zygoptera (Odonata). Systematic Entomology, 39, 68–96.
https://doi.org/10.1111/syen.12035
Ewing, T.E. (1980) Tectonic Evolution of the Pacific Northwest. The Journal of Geology, 88, 619–638.
https://doi.org/10.1086/628551
Fischer, C. (1974) Systematische Stellung der Gattung Sieblosia Handlirsch, 1906 (Zygoptera, Lestinoidea: Sieblosiidae). Odonatologica, 3, 211–220.
Fleck, G., Bechly, G., Martínez-Delclòs, X., Jarzembowski, E.A., Coram, R. & Nel, A. (2003) Phylogeny and classification of the Stenophlebioptera (Odonata: Epiproctophora). Annales de la Société entomologique de France (N.S.): International Journal of Entomology, 39, 55–93.
https://doi.org/10.1080/00379271.2003.10697363
Fleck, G., Bechly, G., Martínez-Delclòs, X., Jarzembowski, E.A. & Nel, A. (2004) A revision of the Upper Jurassic-Lower Cretaceous dragonfly family Tarsophlebiidae, with a discussion on the phylogenetic positions of the Tarsophlebiidae and Sieblosiidae (Insecta, Odonatoptera, Panodonata). Geodiversitas, 26, 33–60.
Fleck, G., Bechly, G., Martínez-Delclòs, X., Jarzembowski, E.A. & Nel, A. (2004) A revision of the Mesozoic dragonfly family Tarsophlebiidae, with a discussion on the phylogenetic positions of the Tarsophlebiidae and Sieblosiidae (Odonatoptera: Panodonata). Geodiversitas, 26, 33–60.
Fleck, G. & Nel, A. (2003) Revision of the Mesozoic family Aeschnidiidae (Odonata: Anisoptera). Zoologica, 153, 1–180.
Fraser, F.C. (1957) A reclassification of the order Odonata, Based on Some New Interpretations of the Venation of the Dragonfly Wing By the late R. J. Tillyard and F. C. Fraser, Revision. Royal Zoological Society of New South Wales, Sydney, 133 pp.
Garrison, R.W., von Ellenrieder, N. & Louton, J.A. (2006) Dragonfly genera of the New World: an illustrated and annotated key to the Anisoptera. Johns Hopkins University Press, Baltimore, Maryland, 368 pp.
Garrison, R.W., von Ellenrieder, N. & Louton, J.A. (2010) Damselfly genera of the New World: an illustrated and annotated key to the Zygoptera. Johns Hopkins University Press, Baltimore, 490 pp.
Garrouste, R. & Nel, A. (2015) New Eocene damselflies and the first Cenozoic damseldragonfly of the isophlebiopteran lineage (Insecta: Odonata). Zootaxa, 4028 (3), 354–366.
https://doi.org/10.11646/zootaxa.4028.3.2
Garrouste, R. & Nel, A. (2019) Alaskan Palaeogene insects: a challenge for a better knowledge of the Beringian ‘route’ (Odonata: Aeshnidae, Dysagrionidae). Journal of Systematic Palaeontology, 17, 1939–1946.
https://doi.org/10.1080/14772019.2019.1572235
Garrouste, R., Wedmann, S., Pouillon, J.-M. & Nel, A. (2017) The oldest ‘amphipterygid’ damselfly of tropical affinities in the Paleocene of Menat (Zygoptera: Eucaloptera). Historical Biology, 29, 818–821.
https://doi.org/10.1080/08912963.2016.1247448
Gentilini, G. & Bagli, L. (2004) Fossil Zygoptera and Anisoptera from the Upper Miocene of Monte Castellaro (Pesaro, Marches, Central Ital). Quaderno di Studi e Notizie di Storio Naturale dello Romagna, 19, 17–44.
Goloboff, P.A. (1993) Estimating character weights during tree search. Cladistics, 9, 83–91.
https://doi.org/10.1111/j.1096-0031.1993.tb00209.x
Goloboff, P.A. & Catalano, S.A. (2016) TNT version 1.5, including a full implementation of phylogenetic morphometrics. Cladistics, 32, 221–238.
https://doi.org/10.1111/cla.12160
Goloboff, P.A., Farris, J.S. & Nixon, K.C. (2008) TNT, a free program for phylogenetic analysis. Cladistics, 24, 774–786.
https://doi.org/10.1111/j.1096-0031.2008.00217.x
Goloboff, P.A., Mattoni, C.I. & Quinteros, A.S. (2006) Continuous characters analyzed as such. Cladistics, 22, 589–601.
https://doi.org/10.1111/j.1096-0031.2006.00122.x
Graham, A. (1999) Late Cretaceous and Cenozoic history of North American vegetation north of Mexico. Oxford University Press, New York, xviii + 350 pp.
https://doi.org/10.1093/oso/9780195113426.001.0001
Greenwood, D.R., Archibald, S.B., Mathewes, R.W. & Moss, P.T. (2005) Fossil biotas from the Okanagan Highlands, southern British Columbia and northern 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, 755 pp.
Gutiérrez, N., Noguera, F.A., Zaragoza-Caballero, S. & Morrone, J.J. (2020) Phylogenetic revision of the floricolous longhorn beetle genus Strangalidium Giesbert (Coleoptera: Cerambycidae). Systematics and Biodiversity, 18, 79–101.
https://doi.org/10.1080/14772000.2020.1729893
Hagen, H. (1848) Die fossilen Libellen Europa’s. Entomologische Zeitung, 9, 6–13.
Hagen, H. (1858) Zwei Libellen aus der Braunkohle von Sieblos. Paleontographica, 5, 121–124.
Handlirsch, A. (1907) Die Fossilen Insekten und die Phylogenie der Rezenten Formen, IV. Tertire Insekten. Ein Handbuch für Paläontologen und Zoologen. Leipzig: Wilhelm Engelmann, pp. 675–1130.
Henriksen, K.A. (1922) Eocene insects of Denmark. Danmarks Geologiske Undersøgelse, 2 række 37, 1–36.
Hobson, K.A., Anderson, R.C., Soto, D.X. & Wassenaar L.I. (2012) Isotopic evidence that dragonflies (Pantala flavescens) migrating through the Maldives come from the northern Indian Subcontinent. PLoS ONE, 7, e52594.
https://doi.org/10.1371/journal.pone.0052594
Huang, D., Azar, D., Cai, C., Maksoud S., Nel, A. & Bechly, G. (2017) Mesomegaloprepidae, a remarkable new damselfly family (Odonata: Zygoptera) from mid-Cretaceous Burmese amber. Cretaceous Research, 73, 1–13.
https://doi.org/10.1016/j.cretres.2017.01.003
Huang, D., Fu, Y. & Nel, A. (2019) The first amber stenophlebiid damsel-dragonfly (Odonata, Epiproctophora, Stenophlebiidae) from mid-Cretaceous amber of northern Myanmar. Cretaceous Research, 94, 40–44.
https://doi.org/10.1016/j.cretres.2018.10.008
Huguet, A., Nel, A., Martinez-Delclos, X. & Bechly, G. (2002) Preliminary phylogenetic analysis of the Protanisoptera (Insecta: Odonatoptera). Geobios, 35, 537–560.
https://doi.org/10.1016/S0016-6995(02)00071-2
International Trust for Zoological Nomenclature (1999) International Code of Zoological Nomencature. Available from: https://www.iczn.org/the-code/the-international-code-of-zoological-nomenclature/ (accessed 27 June 2020)
Jarzembowski, E.A., Martýìnez-Delclòs, X., Bechly, G., Nel, A., Coram, R., & Escuilli, F. (1998) The Mesozoic non-calopterygoid Zygoptera: description of new genera and species from the Lower Cretaceous of England and Brazil and their phylogenetic significance (Odonata, Zygoptera, Coenagrionoidea, Hemiphlebioidea, Lestoidea). Cretaceous Research, 19, 403–444.
https://doi.org/10.1006/cres.1997.0113
Jarzembowski, E.A. & Nel, A. (1996) New fossil dragonflies from the Lower Cretaceous of SE England and the phylogeny of the superfamily Libelluloidea (Insecta: Odonata). Cretaceous Research, 17, 67–85.
https://doi.org/10.1006/cres.1996.0005
Kalkman, V.J. (2006) Key to the dragonflies of Turkey. Brachytron, 10, 3–82.
Kohli, M.K., Ware, J.L. & Bechly, G. (2016) How to date a dragonfly: Fossil calibrations for odonates. Palaeontologia Electronica, 19, 1–14.
Lewis, S.E. (1992) Insects of the Klondike Mountain Formation, Republic, Washington. Washington Geology, 20, 15–19.
Lin, Q.B. (1982) Insecta. In: Palaeontological Atlas of East China. Part 3. Mesozoic and Cenozoic, Geological Publishing Office, Beijing, pp. 148–155.
Lin, Q.-B., Nel, A. & Huang, D.-Y. (2002) Phylogenetic analysis of the Mesozoic dragonfly family Liupanshaniidae (Insecta: Aeshnoptera: Odonata). Cretaceous Research, 23, 439–444.
https://doi.org/10.1006/cres.2002.1013
Maneval, H. (1936) Insectes fossiles des calcaires oligocènes de Ronzon (Le Puy). Annales de la Société Linnéenne de Lyon, 79, 23–37.
https://doi.org/10.3406/linly.1936.14770
Martínez-Delclòs, X. & Martinell, J. (1993) Insect taphonomy experiments. Their application to the Cretaceous outcrops of lithographic limestones from Spain. Kaupia. Darmstädter Beiträge zur Naturgeschichte, 2, 133–144.
Mathewes, R.W., Greenwood, D.G. & Archibald, S.B. (2016) Paleoevironment of the Quilchena Flora, British Columbia, during the Early Eocene climatic optimum. Canadian Journal of Earth Sciences, 53, 574–590.
https://doi.org/10.1139/cjes-2015-0163
Mongiardino Koch, N.M., Soto, I.M. & Ramírez, M.J. (2015) First phylogenetic analysis of the family Neriidae (Diptera), with a study on the issue of scaling continuous characters. Cladistics, 31, 142–165.
https://doi.org/10.1111/cla.12084
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
Nel, A. (1986) Révision du genre cénozoïque Stenolestes Scudder, 1895; description de deux especes nouvelles (Insecta, Odonata, Lestidae). Bulletin du Muséum national d’histoire naturelle, Paris, (4) (C), 8, 447–461.
Nel, A. (1988) Parazacallitinae, nouvelle sous-famille et premier Epallagidae’ de l’Oligocène européen (Odonata, Zygoptera). Bulletin du Muséum National d’Histoire Naturelle, Paris, 10, 175–179.
Nel, A. (1991) Description de quelques Sieblosiidae fossiles nouveaux (Odonata, Zygoptera, Lestoidea). Nouvelle Revue d’Entomologie, New Series, 8, 367–375.
Nel, A. & Arillo, A. (2006) The first Baltic amber dysagrionine damselfly (Odonata: Zygoptera: Thaumatoneuridae: Dysagrioninae), Annales de la Société entomologique de France, New Series, International Journal of Entomology, 42, 179–182.
https://doi.org/10.1080/00379271.2006.10700621
Nel, A., Arillo, A. & Martínez-Delclòs, X. (1996) New fossil Odonata (Insecta) from the Upper Miocene of France and Spain (Anisoptera and Zygoptera). Neues Jahrbuch für Geologie und Paläontologie—Abhandlungen, 199, 167–219.
https://doi.org/10.1127/njgpa/199/1996/167
Nel, A., Bechly, G., Jarzembowski, E.A. & Martínez-Delclòs, X. (1998) A Revision of the Fossil Petalurid Dragonflies (Insecta: Odonata: Anisoptera: Petalurida). Paleontologica Lombarda, 10, 3–69.
Nel, A., Bethoux, O., Bechly, G., Martínez-Delclòs, X. & Papier, F. (2001) The Permo-Triassic Odonatoptera of the ‘Protodonate’ Grade (Insecta: Odonatoptera). Annales de la Sociét Entomologique de France, 37, 501–525.
Nel, A. & Escuillié, F. (1993) Découverte d’un nouveau genre de Sieblosiidae dans le Miocène supérieur de l’Ardèche (France) (Odonata, Zygoptera, Lestoidea, Sieblosiidae). Nouvelle Revue d’Entomologie, New Series, 10, 233–242.
Nel, A. & Fleck, G. (2012) A new genus and species of Sieblosiidae from the middle Miocene of Germany (Odonata: Epiproctophora). Zootaxa, 3582 (1), 64–68.
https://doi.org/10.11646/zootaxa.3582.1.7
Nel, A. & Fleck, G. (2014) Dragonflies and damselflies (Insecta: Odonata) from the Late Eocene of the Isle of Wight. Earth and Environmental Science Transactions of the Royal Society of Edinburgh, 104, 283–306.
https://doi.org/10.1017/S175569101400005X
Nel, A., Fleck, G., Garrouste, R., Gand, G., Lapeyrie, J., Bybee, S.M. & Prokop, J. (2009) Revision of Permo-Carboniferous griffenflies (Insecta: Odonatoptera: Meganisoptera) based upon new species and redescription of selected poorly known taxa from Eurasia. Palaeontographica (A), 289, 89–121.
https://doi.org/10.1127/pala/289/2009/89
Nel, A., Petrulevicius, J.F., Gentilini, G. & Martínez-Delclòs, X. (2005c) Un nouvel Odonate du Miocène d’Italie (Odon.). Bulletin de la Société entomologique de France, 110, 188.
Nel, A., Martinez-Delclos, X., Paicheler, J.-C. & Henrotay, M. (1993) Les “Anisozygoptera” fossiles: phylogénie et classification (Odonata). Martinia, 3, 1–311.
Nel, A., Martínez-Delclòs, X., Papier, F. & Oudard, J. (1997) New Tertiary fossil Odonata from France. (Sieblosiidae, Lestidae, Coenagrioniidae, Megapodagrionidae, Libellulidae). Deutsche Entomologische Zeitschrift, 44, 231–258.
https://doi.org/10.1002/mmnd.19970440210
Nel, A. & Paicheler, J.-C. (1992) Les Odonata fossiles: Etat actuel des connaissances. Huitième partie: Les Calopterygoidea fossiles (Odonata, Zygoptera). Bulletin de la Société Entomologique de France, 97, 381–396.
Nel, A. & Paicheler, J.-C. (1994) Les Lestoidea (Odonata, Zygoptera) fossiles: Un inventaire critique. Annales de Paléontologie, 80, 1–59.
Nel, A. & Papazian, M. (1986) Sur une nouvelle espece d’Odonate fossile du Stampien de Cereste (Luberon). Nouvelle Revue d’Entomologie, New Series, 3, 227–233.
Nel, A., Petrulevičius, J.F., Gentilini, G. & Martínez-Delclòs, X. (2005a) Phylogenetic analysis of the Cenozoic family Sieblosiidae (Insecta: Odonata), with description of new taxa from Russia, Italy and France. Geobios, 38, 219–233.
https://doi.org/10.1016/j.geobios.2003.10.007
Nel, A., Petrulevičius J.F. & Jarzembowsk, E.A. (2005b) New fossil Odonata from the European Cenozoic (Insecta: Odonata: Thaumatoneuridae, Aeshnidae, ?Idionychidae, Libellulidae). Neues Jahrbuch für Geologie und Paläontologie, Abhandlungen, 235, 343–380.
Nel, A., Simov, N., Bozukov, V. & Marinov, M. (2016) New dragonflies and damselflies from Middle Miocene deposits in SW Bulgaria (Insecta: Odonata). Palaeontologia Electronica, 19.3.35A, 1–13.
Parins-Fukuchi, C. (2018) Use of continuous traits can improve morphological phylogenetics. Systematic Biology, 67, 328–339.
https://doi.org/10.1093/sysbio/syx072
Pessacq, P. (2008) Phylogeny of Neotropical Protoneuridae (Odonata: Zygoptera) and a preliminary study of their relationship with related families. Systematic Entomology, 33, 511–528.
https://doi.org/10.1111/j.1365-3113.2007.00414.x
Petrulevičius, J.F. (2017) First Frenguelliidae (Insecta: Odonata) from the middle Eocene of Río Pichileufú, Patagonia, Argentina. Arquivos Entomolóxicos, 18, 367–374.
Petrulevičius, J.F. (2019) A new Frenguelliidae (Insecta: Odonata) from the Eocene of Arroyo Chacay, Patagonia, Argentina. Palaeoentomology, 002, 591–595.
https://doi.org/10.11646/palaeoentomology.2.6.9
Petrulevičius, J.F. & Gutiérrez, P.R. (2016) New basal Odonatoptera (Insecta) from the lower Carboniferous (Serpukhovian) of Argentina. Arquivos Entomolóxicos, 16, 341–358.
Petrulevičius, J.F. & Nel, A. (2003) Frenguelliidae, a new family of dragonflies from the earliest Eocene of Argentina (Insecta: Odonata): phylogenetic relationships within Odonata. Journal of Natural History, 37, 2909–2917.
https://doi.org/10.1080/0022293021000007543
Petrulevičius, J.F. & Nel, A. (2007) Enigmatic and little known Odonata (Insecta) from the Paleogene of Patagonia and Northwest Argentina. Annales de la Société entomologique de France, New Series, 43, 341–347.
https://doi.org/10.1080/00379271.2007.10697530
Petrulevičius, J.F. & Nel A. (2013) A new Frenguelliidae (Insecta: Odonata) from the early Eocene of Laguna del Hunco, Patagonia, Argentina. Zootaxa, 3616 (6), 597–600.
https://doi.org/10.11646/zootaxa.3616.6.6
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. Alevesia, 1, 15–25.
Petrulevičius, J.F., Wappler, T., Wedmann, S., Rust, J. & Nel, A. (2008) New Megapodagrionid Damselflies (Odonata: Zygoptera) from the Paleogene of Europe. Journal of Paleontology, 82, 1173–1181.
https://doi.org/10.1666/07-091.1
Rasnitsyn, A.P. & Pritykina, L.N. (2002) Superorder Libellulidea Laicharting, 1781. Order Odonata Fabricius, 1792. The Dragonflies. In: Rasnitsyn, A.P. & Quicke, D.L.J. (Eds.), History of Insects. Kluwer Academic Publishers, London, pp. 97–104.
Rehn, A.C. (2003) Phylogenetic analysis of higher-level relationships of Odonata. Systematic Entomology, 28, 181–239.
https://doi.org/10.1046/j.1365-3113.2003.00210.x
Riek, E.F. & Kukalová-Peck, J. (1984) A new interpretation of dragonfly wing venation based upon Early Upper Carboniferous fossils from Argentina (Insecta: Odonatoidea) and basic character states in pterygote wings. Canadian Journal of Zoology, 62, 1150–1166.
https://doi.org/10.1139/z84-166
Riou, B. & Nel, A. (1995) Nouveaux Odonates fossiles du Miocène supérieur de l’Ardèche. (Odonata: Sieblosiidae, Lestidae, Aeshnidae, Libellulidae, Corduliidae). E.P.H.E., Biologie et Evolution des Insectes, Paris, 7/8, 125–144.
Rouse, G.E., Hopkins, W.S. Jr. & Piel, K.M. (1971) 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. Postdoctoral thesis, Biologische Fakultät der Georg-August-Universität Göttingen, Göttingen, Lower Saxony, 482 pp.
Rust, J, Petrulevičius, J.F. & Nel, A. (2008) The first damselflies from the lowermost Eocene of Denmark, with a description of a new subfamily (Odonata, Zygoptera, Dysagrionidae). Palaeontology, 51, 709–713.
https://doi.org/10.1111/j.1475-4983.2008.00780.x
Scudder, S.H. (1877) The insects of the Tertiary beds at Quesnel (British Columbia). Report of progress of the Geological Survey of Canada 1875–1876, 1877, 266–280.
Scudder, S.H. (1878) An Account of some insects of unusual interest from the Tertiary rocks of Colorado and Wyoming. Bulletin of the United States Geological and Geographical Survey of the Territories, 4, 519–543.
Scudder, S.H. (1885) 4. Classe Insecta. Insecten. Handbuch der Palaeontologie, I Abtheilung. Palaeozoologie, 2, 747–831.
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.
https://doi.org/10.5962/bhl.title.44698
Scudder, S.H. (1895) The Miocene insect fauna of Oenegin, Baden. Geological Magazine, 2, 116–122.
https://doi.org/10.1017/S0016756800005963
Serrano-Meneses, M.A., Córdoba-Aguilar, A., Azpilicueta-Amorín, M., Gonzáles-Soriano, E. & Székely, T. (2008) Sexual selection, sexual size dimorphism and Rensch’s rule in Odonata. Journal of Evolutionary Biology, 21, 1259–1273.
https://doi.org/10.1111/j.1420-9101.2008.01567.x
Statz, G. (1930) Drei neue Insektenarten aus dem Tertiär von Rott am Siebengebirge. Wissenschaftliche Mitteilungen des Vereins für Natur und Heimatkunde in Köln, 1, 10–14.
Théobald, N. (1937) Les Insectes fossiles des terrains oligocènes de France. Thesis, Université de Nancy, Nancy, 467, pp, 29 pls.
Tillyard, R.J. & Fraser, F.C. (1939) A reclassification of the order Odonata based on some new interpretations of the dragonfly wing by R.J. Tillyard, continuation thereof by F.C. Fraser, Part II, the suborder Zygoptera (continued), Protanisoptera. Australian Zoologist, 9, 195–221.
Tillyard, R.J. & Fraser, F.C. (1940) A reclassification of the order Odonata based on some new interpretations of the dragonfly wing by R.J. Tillyard, continuation thereof by F.C. Fraser, Part III, suborder Anisozygoptera. Australian Zoologist, 9, 359–396.
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
US National Parks Service (2015) Florissant National Monument fossil database. Available from: https://www.nps.gov/flfo/learn/education/introduction-to-the-fossil-database.htm (accessed 27 June 2020)
Wagner, T., Neinhuis, C. & Barthlott, W. (1996) Wettability and contaminability of insect wings as a function of their surface sculptures. Acta Zoologica, 77, 213–225.
https://doi.org/10.1111/j.1463-6395.1996.tb01265.x
Wedmann, S., Uhl, D., Lehmann, T., Garrouste, R. Nel, A., Gomez, B., Smith, K. & Schaal, S.F.K. (2018) The Konservat-Lagerstätte Menat (Paleocene; France) – an overview and new insights. Geologica Acta, 16, 189–213.
Wehr, W.C. & Barksdale, L.L. (1996) A Checklist of Fossil Insects from Republic, Washington. Washington Geology, 24, 29.
Westfall, M.J., May, M.L. & Needham, J.G. (2014) Dragonflies of North America: The Odonata (Anisoptera) Fauna of Canada, the Continental United States, Northern Mexico and the Greater Antilles. Scientific Publishers, Gainesville, Florida, xiv + 648 pp.
Wilf, P., Cúneo, N.R. Johnson, K.R., Hicks, J.F., Wing, S.L. & Obradovich, J.D. (2003) High plant diversity in Eocene South America: evidence from Patagonia. Science, 300, 122–125.
https://doi.org/10.1126/science.1080475
Wilson, M.V.H. (1977) 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
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]
Wolfe, J.A. & Wehr, W.C. (1987) Middle Eocene Dicotyledonous plants from Republic, northeast Washington. United States Geological Survey Bulletin, 1597, 1–25, 16 pls.
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
Zachos, J., Pagani, M., Sloan, L., Thomas, E. & Billups, K. (2001) Trends, rhythms, and aberrations in global climate 65 Ma to present. Science, 292, 686–693.
https://doi.org/10.1126/science.1059412
Zheng, D. & Jarzembowski, E.A. (2020) A brief review of Odonata in mid-Cretaceous Burmese amber. International Journal of Odonatology, 23, 13–21.
https://doi.org/10.1080/13887890.2019.1688499
Zessin, W. (2011) Neue Insekten aus dem Moler (Paläozän/Eozän) von Dänemark Teil 1 (Odonata: Epallagidae, Megapodagrioniidae). Virgo, Mitteilungsblatt des Entomologischen Vereins Mecklenburg, 14, 63–73.
Zhang, J.-F. (1992) Congqingia rhora gen. nov., spec. nov. – a new dragonfly from the Upper Jurassic of eastern China (Anisozygoptera, Congquingiidae fam. nov.). Odonatologica, 21, 375–383.
Zheng, D., Chang, S.-C., Nel, A., Jarzembowski, E.A., Zhuo, D. & Wang, B. (2017) Electrodysagrion lini gen. et sp. nov., the oldest Dysagrionini (Odonata: Zygoptera: Dysagrionidae) from mid-Cretaceous Burmese amber. Cretaceous Research, 77, 44–48.
https://doi.org/10.1016/j.cretres.2017.05.008
Zheng, D, Chang, S.-C. & Wang, B. (2018) A new dysagrionid damselfly (Odonata: Zygoptera: Palaeodysagrion) from mid-Cretaceous Burmese amber. Alcheringa, 42, 300–304.
https://doi.org/10.1080/03115518.2017.1381991
Zheng, D. Wang, B. Jarzembowski, E.A. Chang, S.-C. & Nel, A. (2016a) Burmadysagrioninae, a new subfamily (Odonata: Zygoptera: Dysagrionidae) from mid-Cretaceous Burmese amber. Cretaceous Research, 67, 126–132.
https://doi.org/10.1016/j.cretres.2016.07.006
Zheng, D., Zhang, Q., Nel, A., Jarzembowski, E.A. & Wang, B. (2019) Electrodysagrion neli sp. nov., the second Cretaceous dysagrionine damselfly (Odonata: Zygoptera: Dysagrionidae) from Kachin amber, northern Myanmar. Palaeoentomology, 2, 556–559.
https://doi.org/10.11646/palaeoentomology.2.6.2
Zheng, D., Zhang, Q., Nel, A., Jarzembowski, E.A., Zhou, Z., Chang, S.-C. & Wang, B. (2016b) New damselflies (Odonata: Zygoptera: Hemiphlebiidae, Dysagrionidae) from mid-Cretaceous Burmese amber. Alcheringa, 41, 12–21.
https://doi.org/10.1080/03115518.2016.1164402
Zhou, J.-B., Han, W., Song, M.-C. & Li, L. (2020) Zircon U-Pb ages of the cetaceous [sic] sedimentary rocks in the Laiyang Basin, eastern China and their tectonic implications. Journal of Asian Earth Sciences. [early issue]
https://doi.org/10.1016/j.jseaes.2019.103956
Zittel, K.A. (1885) Classe Insecta. Insecten (Scudder, S.H. Ed.) In: Handbuch der Palaeontologie. Vol. 1. Abtheilung Palaeozoologie, Band 2, Mollusca and Arthropoda, R. Oldenbourg, München and Liepzig, pp. 747–831.