Skip to main content Skip to main navigation menu Skip to site footer
Type: Monograph
Published: 2017-05-25
Page range: 1–63
Abstract views: 410
PDF downloaded: 17

Fossil butterflies, calibration points and the molecular clock (Lepidoptera: Papilionoidea)

Naturalis Biodiversity Center, P.O. Box 9517, 2300RA Leiden, The Netherlands
biogeography calibration points evolutionary history morphology molecular clock phylogeny

Abstract

Fossil butterflies are extremely rare. Yet, they are the only direct evidence of the first appearance of particular characters and as such, they are crucial for calibrating a molecular clock, from which divergence ages are estimated. In turn, these estimates, in combination with paleogeographic information, are most important in paleobiogeographic considerations. The key issue here is the correct allocation of fossils on the phylogenetic tree from which the molecular clock is calibrated.

The allocation of a fossil on a tree should be based on an apomorphic character found in a tree based on extant species, similar to the allocation of a new extant species. In practice, the latter is not done, at least not explicitly, on the basis of apomorphy, but rather on overall similarity or on a phylogenetic analysis, which is not possible for most butterfly fossils since they usually are very fragmentary. Characters most often preserved are in the venation of the wings. Therefore, special attention is given to possible apomorphies in venational characters in extant butterflies. For estimation of divergence times, not only the correct allocation of the fossil on the tree is important, but also the tree itself influences the outcome as well as the correct determination of the age of the fossil. These three aspects are discussed.

        All known butterfly fossils, consisting of 49 taxa, are critically reviewed and their relationship to extant taxa is discussed as an aid for correctly calibrating a molecular clock for papilionoid Lepidoptera. In this context some aspects of age estimation and biogeographic conclusions are briefly mentioned in review. Specific information has been summarized in four appendices.

References

  1. Ackery, P.R., de Jong, R. & Vane-Wright, R.I. (1999) The butterflies: Hedyloidea, Hesperioidea and Papilionoidea. In: Kristensen, N.P. (Ed.), Lepidoptera, Moths and Butterflies. 1. Evolution, Systematics, and Biogeography. Handbook of Zoology. 4 (35). Lepidoptera. De Gruyter, Berlin, pp. 263–300.

    Ansorge, J. (2015) Günther Wangrins Tagfalter-Versteinerung in einer Stettiner Kugel—die Geschichte einer Fälschung. Geschiebekunde Aktuell, 31, 98–103.

    Bachofen-Echt, J. (1949) Der Bernstein und seine Einschlüsse. Springer Verlag, Wien, 204 pp.
    https://doi.org/10.1007/978-3-7091-2303-4

    Bascombe, M.J., Johnston, G. & Bascombe, F.S. (1999) The Butterflies of Hong Kong. Academic Press, London, ix + 422 pp.

    Behrensmeyer, A.K., Damuth, J.D., DiMichele, W.A., Potts, R., Sues, H.-D. & Wing, S.L. (1992). Terrestrial Ecosystems through Time. University of Chicago Press, Chicago, xix + 568 pp.

    Benton, M., Donoghue, P.C.J. & Asher, R.J. (2009) Calibrating and constraining molecular clocks. In: Hedges, S.B. & Kumar, S. (Eds.), The Timetree of Life. Oxford University Press, Oxford, pp. 35–86.

    Berggren, W.A. & Prothero, D.R. (1993) Eocene-Oligocene climatic and biotic evolution: an overview. In: Prothero, D.R. & Berggren, W.A. (Eds.), Eocene-Oligocene Climatic and Biotic Evolution. Princeton University Press, Princeton, pp. 1–28.

    Boisduval, J.A. (1841) [Cyllo sepulta]. Annales de la Société entomologique de France, 9, 371–374.

    Braby, M.F., Trueman, J.W.H. & Eastwood, R. (2005) When and where did troidine butterflies (Lepidoptera: Papilionidae) evolve? Phylogenetic and biogeographic evidence suggests an origin in remnant Gondwana in the Late Cretaceous. Invertebrate Systematics, 19, 113–143.
    https://doi.org/10.1071/IS04020

    Braby, M.F., Vila, R. & Pierce, N.E. (2006) Molecular phylogeny and systematics of the Pieridae (Lepidoptera: Papilionoidea): higher classification and biogeography. Zoological Journal Linnean Society, 147, 239–275.
    https://doi.org/10.1111/j.1096-3642.2006.00218.x

    Brakefield, P.M., Gates, J., Keys, D., Kesbeke, F., Wijngaarden, P.J., Monteiro, A., French, V. & Carroll, S.B. (1996) Development, plasticity and evolution of butterfly eyespot patterns. Nature, 384, 236–242.
    https://doi.org/10.1038/384236a0

    Branscheid, F. (1968) Ein weiterer Schmetterlingsflügel aus dem Pliozän von Willershausen. Beihefte Berichte Naturhistorisches Gesellschaft Hannover, 6, 41–42.

    Branscheid, F. (1969) Funde von Tagfaltern (Rhopalocera, Lepidopt.) im Pliozän von Willershausen. Berichte Naturhistorisches Gesellschaft Hannover, 113, 101–106.

    Bricoux, F. (1975) Le Genre Zerynthia Ochsenheimer, 1816 (Lepidoptères Papilionidae) en Provence. Sciences naturelles, Paris, 84 pp.

    Brito, I.M. & Ribeiro, F.A.M. (1975) Ocorrência de Lepidoptera nos Folhelhos de Tremembé e algumas considerações sobre a Bacia Geológica do Paraíba, Estado de São Paulo. Anais da Academia Brasiliera Ciências, 47, 106–111.

    Brower, A.V.Z. & Egan, M.G. (1997) Cladistic analysis of Heliconius butterflies and relatives (Nymphalidae: Heliconiiti): A revised phylogenetic position or Eueides based on sequences from mtDNA and a nuclear gene. Proceedings of the Royal Society B: Biological Sciences, 264, 969–977.
    https://doi.org/10.1098/rspb.1997.0134

    Brower, A.V.Z. & Vane-Wright, R.I. (2011) Anomalous areas and awkward ages: concerns about over-reliance on model-based biogeographical and temporal inferences. Systematic Entomology, 36, 601–603.
    https://doi.org/10.1111/j.1365-3113.2011.00586.x

    Brower, A.V.Z., Freitas, A.V.L., Lee, M.-M., Silva-Brandão, K.L., Whinnet, A. & Willmott, K.R. (2006) Phylogenetic relationships among the Ithomiini (Lepidoptera: Nymphalidae) inferred from one mitochondrial and two nuclear gene regions. Systematic Entomology, 31, 288–301.
    https://doi.org/10.1111/j.1365-3113.2006.00321.x

    Brower, A.V.Z., Wahlberg, N., Ogawa, J.R., Boppré, M. & Vane-Wright, R.I. (2010) Phylogenetic relationships among genera of danaine butterflies (Lepidoptera: Nymphalidae) as implied by morphology and DNA sequences. Systematics and Biodiversity, 8, 75–89.
    https://doi.org/10.1080/14772001003626814

    Brower, A.V.Z., Willmott, K.R., Silva-Brandão, K.L., Garzón-Orduña, I.J.& Freitas, A.V.L. (2014) Phylogenetic relationships of ithomiine butterflies (Lepidoptera: Nymphalidae: Danainae) as implied by combined morphological and molecular data. Systematics and Biodiversity, 12, 133–147.
    https://doi.org/10.1080/14772000.2014.899650

    Brown, F.M. & Heineman, B. (1972) Jamaica and its butterflies. E.W. Classy Ltd, London, 478 pp.

    Brown, F.M. (1976) †Oligodonta florissantensis, gen. n., sp. nov. (Lepidoptera: Pieridae). Bulletin Allyn Museum, 37, 1–4.

    Brown, F.M. (1978) The origins of the West Indian butterfly fauna. Academy of Natural Sciences of Philadelphia, Special Publication, 13, 5–30.

    Bryk, F. (1913) Über eine neue Einteilung der Papilionidae unter Berücksichtigung des Flügelgeäders. Archiv für Naturgeschichte, 79 (A2), 116–121.

    Bryk, F. (1914) Parnassius apollo und sein Formenkreis. Verlagsbuchhandlung R. Stricker, Berlin, 148 pp.

    Bryk, F. (1934) Baroniidae, Teinopalpidae, Parnassiidae pars 1. In: Das Tierreich 64. De Gruyter, Berlin, Leipzig, xxiii + 790 pp.

    Butler, A.G. (1873) On fossil butterflies. Lepidoptera Exotica, 15, 126–128.

    Butler, A.G. (1889) Description of a new genus of fossil moths belonging to the geometrid family Euschemidae. Proceedings zoological Society London, 1889, 292–297.

    Butler, A.G. (1894) On two collections of Lepidoptera sent by Mr. H.H. Johnston from British Central Africa. Proceedings of the Zoological Society of London, 1893 (4), 643–684.

    Charpentier, T. de (1843) Über einige fossile Insecten aus Radoboj in Croatia. Verhandlungen der Kaiserlichen Leopoldinisch–Carolinischen Akademie der Naturforscher, 12, 399–410, pls. 20–23.

    CoBabe, E.A., Chamberlain, K.R., Ivie, M.A. & Giersch, J.J. (2002) A new insect and plant Lagerstätte from a Tertiary lake deposit along the Canyon Fery Reservoir, Southwestern Montana. Rocky Mountain Geology, 37, 13–30.
    https://doi.org/10.2113/gsrocky.37.1.13

    Cockerell, T.D.A. (1907) A fossil butterfly of the genus Chlorippe. Canadian Entomologist, 39, 361–363.
    https://doi.org/10.4039/Ent39361-11

    Cockerell, T.D.A. (1908) Fossil insects from Florissant, Colorado. Bulletin of the American Museum of Natural History, 24, 59–69.

    Cockerell, T.D.A. (1909) 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. (1913) Some fossil insects from Florissant, Colorado. Proceedings of the United States National Museum, 44, 341–346.
    https://doi.org/10.5479/si.00963801.44-1955.341

    Common, I.F.B. (1973) A new family of Dacnonypha (Lepidoptera) based on three new species from southern Australia, with notes on the Agathiphagidae. Journal of the Australian Entomological Society, 12, 11–23.
    https://doi.org/10.1111/j.1440-6055.1973.tb02150.x

    Comstock, J.H. (1918) The Wings of Insects. Comstock, Ithaca, N.Y., xvii + 430 pp.

    Comstock, W.P. (1961) Butterflies of the American Tropics: the Genus Anaea, Lepidoptera Nymphalidae. American Museum of Natural History, New York, 214 pp.

    Condamine, F.L., Sperling, F.A.H., Wahlberg, N., Rasplus, J.-Y. & Kergoat, G. (2012) What causes latitudunal gradients in species diversity? Evolutionary processes and ecological constraints on swallowtail biodiversity. Ecology Letters, 15, 267–277.
    https://doi.org/10.1111/j.1461-0248.2011.01737.x

    Cottrell, C.B. (1984) A phytophagy in butterflies: its relationship to myrmecophily. Zoological Journal of the Linnean Society, 80, 1–57.
    https://doi.org/10.1111/j.1096-3642.1984.tb02318.x

    Crowley, T.J. & North, G.R. (1991) Paleoclimatology. Oxford University Press, Oxford, 326 pp.

    Daudet, H. (1876) Descriptions d'une chenille fossile trouvée dans le calcaire d'Aix (Provence). Revue et Magasin de Zoologie, 1876, 415–424.

    de Jong, R. (1980) Some tools for evolutionary and phylogenetic studies. Zeitschrift für zoologische Systematik und Evolutions-forschung, 18, 1–23.
    https://doi.org/10.1111/j.1439-0469.1980.tb00724.x

    de Jong, R. (2007) Estimating time and space in the evolution of the Lepidoptera. Tijdschrift voor Entomologie, 150, 319–346.
    https://doi.org/10.1163/22119434-900000233

    de Jong, R. (2016) Reconstructing a 55-million-year old butterfly (Lepidoptera, Hesperiidae). European Journal of Entomology, 113, 423–428.
    https://doi.org/10.14411/eje.2016.055

    de Jong, R., Vane-Wright, R.I. & Ackery, P.R. (1996) The higher classification of butterflies: problems and prospects. Entomologica Scandinavica, 27, 65–101.
    https://doi.org/10.1163/187631296X00205

    Demoulin, G. (1975) A propos de deux Papilionidae méditerranéens fossiles. Bulletin Cercle des Lépidopteristes de Belgique, 4, 96–101.

    DeVries, P.J. & Poinar, G.O. (1997) Ancient butterfly-ant symbiosis: direct evidence from Dominican amber. Proceedings of the Royal Society of London B, 264, 1137–1140.
    https://doi.org/10.1098/rspb.1997.0157

    DeVries, P.J. (1987) The Butterflies of Costa Rica`and their natural history: Papilionidae, Pieridae, and Nymphalidae. Princeton University Press, Princeton, xxv + 288 pp.

    DeVries, P.J. (1997) The Butterflies of Costa Rica and their natural history. Vol. II. Riodinidae. Princeton University Press, Princeton, xxii + 327 pp.

    Drummond, A.J. & Rambaut, A. (2007) BEAST: Bayesian evolutionary analysis by sampling trees. BMC Evolutionary Biology, 7, 214.
    https://doi.org/10.1186/1471-2148-7-214

    Durden, C.J. & Rose, H. (1978) Butterflies from the Middle Miocene: the earliest occurrence of fossil Papilionoidea (Lepidoptera). Pearce-Sellards Series, 29, 1–25.

    Eliot, J.N. (1973) The higher classification of the Lycaenidae (Lepidoptera): a tentative arrangement. Bulletin of the British Museum (Natural History), Entomology, 28, 373–505, pls. 1–6.
    https://doi.org/10.5962/bhl.part.11171

    Emmel, T.C., Minno, M.C. & Drummond, B.A. (1992) Florissant butterflies. A guide to the fossil and present-day species of central Colorado. Stanford University Press, Stanford, x + 118 pp.

    Eskov, K. Yu. (2002) Geographical history of insects. In: Rasnitsyn, A.P. & Quicke, D.L.J. (Eds.), History of Insects. Kluwer Academic Publishers, Dordrecht, pp. 427–435.

    Espeland, M., Hall, J.P.W., DeVries, P.J., Lees, D.C., Cornwall, M., Hsu, Y-F., Wu, L-W., Campbell, D.L., Talavera, G., Vila, R., Salzman, S., Ruehr, S., Lohman, D.J. & Pierce, N. (2015) Ancient Neotropical origin and recent recolonization: phylogeny, biogeography and diversification of the Riodinidae (Lepidoptera: Papilionoidea). Molecular Phylogenetics and Evolution, 93, 296–306.
    https://doi.org/10.1016/j.ympev.2015.08.006

    Evers, J. (1907) Copal-Schmetterlinge. Entomologisches Jahrbuch, 1907, 129–132.

    Forbes, A.T.M. (1932) How old are the Lepidoptera? American Naturalist, 66, 452–460.
    https://doi.org/10.1086/280451

    Forey, P.L. (1992) Fossils and cladistic analysis. In: Forey, P.L., Humphries, C.J., Kitching, I.J., Scotland, R.W., Siebert, D.J. & Williams, D.M. (Eds.), Cladistics, a Practical Course in Systematics. Clarendon Press, Oxford, pp. 124–136.

    Fox, R.M. (1948) The scarcest of fossils. Carnegie Magazine, 22, 51–52.

    Freitas, A.V.L. & Brown, K.S. (2004) Phylogeny of the Nymphalidae (Lepidoptera). Systematic Biology, 53, 363–383.
    https://doi.org/10.1080/10635150490445670

    Fric, Z., Wahlberg, N., Pech, P. & Zrzavy, J. (2007) Phylogeny and classification of the Phengaris-Maculinea clade (Lepidoptera: Lycaenidae): total evidence and phylogenetic species concepts. Systematic Entomology, 32, 558–567.
    https://doi.org/10.1111/j.1365-3113.2007.00387.x

    Fujiyama, I. (1968) A Pleistocene fossil Papilio from Shiobara, Japan. Bulletin of the National Science Museum, Tokyo, 11, 85–95.

    Fujiyama, I. (1983) A Pleistocene nymphalid butterfly from Shiobara, Japan. Bulletin of the National Science Museum, Tokyo (C), 9, 121–128.

    Gaede, M. (1931) Satyridae. Lepidopterorum Catalogus, 29, 1–759.

    Garzón-Orduña, I.J., Silva-Brandão, K.L., Willmott, K.R., Freitas, A.V.L. & Brower, A.V.Z. (2015) Incompatible ages for clearwing butterflies based on alternative decondary calibrations. Systematic Biology, 64, 752–767.
    https://doi.org/10.1093/sysbio/syv032

    Gingerich, P.D. (1979) Paleontology, phylogeny, and classification: an example from the mammalian fossil record. Systematic Zoology, 28, 451–464.
    https://doi.org/10.2307/2412560

    Grande, L. (2013) The Lost World of Fossil Lake: Snapshots from Deep Time. University of Chicago Press, Chicago, 432 pp.
    https://doi.org/10.7208/chicago/9780226922980.001.0001

    Gravenhorst, J.L.C. (1835) Bericht der entomologischen Section. Schlesische Gesellschaft für Vaterländische Kultur Jahresbericht, 1834, 88–95.

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

    Hall, J.P.W. (2003) Phylogenetic reassessment of the five forewing radial-veined tribes of Riodininae (Lepidoptera: Riodinidae). Systematic Entomology, 28, 23–37.
    https://doi.org/10.1046/j.1365-3113.2003.00196.x

    Hall, J.P.W., Robbins, R.K. & Harvey, D.J. (2004) Extinction and biogeography in the Caribbean: new evidence from a fossil riodinid butterfly in Dominican amber. Proceedings of the Royal Society of London B, 271, 797–801.
    https://doi.org/10.1098/rspb.2004.2691

    Hammond, P.C. & Poinar Jr., G. (1998) A larval brush-footed butterfly (Lepidoptera: Nymphalidae) in Dominican amber, with a summary of fossil Nymphalidae. Entomologica Scandinavica, 29, 275–279.
    https://doi.org/10.1163/187631298X00104

    Hancock, D.L. (1983) Classification of the Papilionidae (Lepidoptera): a phylogenetic approach. Smithersia, 2, 1–48.

    Handlirsch, A. (1908) Die fossilen Insekten und die Phylogenie der rezenten Formen. Wilhelm Engel, Leipzig, ix + 1430 pp.

    Handlirsch, A. (1925) Handbuch der Entomologie. Bd. III. Fischer, Jena, xiii + 1201 pp.

    Harvey, D.J. (1987) The higher classification of the Riodinidae, Lepidoptera. Ph.D. thesis, University of Texas, Austin, 215 pp.

    Heer, O. (1849) Die Insektenfauna der Tertiärgebilde von Oeningen und von Radoboj in Croatien. Vol. 2. Wilhelm Engelsmann, Leipzig, 264 pp.

    Heikkilä, M., Kaila, L., Mutanen, M., Peña, C. & Wahlberg, N. (2011) Cretaceous origin and repeated Tertiary diversification of the redefined butterflies. Proceedings of the Royal Society B, 279, 1093–1099.
    https://doi.org/10.1098/rspb.2011.1430

    Hemming, F. (1967) The generic names of the butterflies and their type-species (Lepidoptera: Rhopalocera). Bulletin of the British Museum (Natural History), Entomology, 9 (Supplement), 1–509.

    Hennig, W. (1966a) Phylogenetic Systematics. University of Illinois Press, Urbana, 263 pp.

    Hennig, W. (1966b) Fannia scalaris Fabricius, eine rezente Art im baltischen Bernstein? (Diptera: Muscidae). Stuttgarter Beiträge zur Naturkunde, 150, 1–12.

    Hennig, W. (1969) Die Stammesgeschichte der Insekten. Waldemar Kramer, Frankfurt, 436 pp.

    Hennig, W. (1981) Insect phylogeny. John Wiley & Sons, Chichester, etc., xxii + 514 pp.

    Henrotay, M. (1986) Découverte d’un nouveau gisement de Lépidoptères, d’autres insectes et d’Araignés fossiles à Dauphin (Hte Provence, France). Linneana belgica, 10, 266–279.

    Herrich-Schäffer, G.A.W. (1843–1856) Systematische Bearbeitung der Schmetterlinge von Europa. Vols. 1–6. In Commission bei G. J. Manz, Regensburg, 2000 pp.

    Hillis, D.M., Moritz, C. & Mable, B. (Eds.) (1996) Molecular Systematics. 2nd Edition. Sinauer Associates, Inc., Sunderland, Massachusetts, xvi + 655 pp.

    Howe, W.H. (1975) The Butterflies of North America. Doubleday & Comp, Inc., Garden City, New York, xiii + 633 pp., 97 pls.

    Igarashi, S. & Harada, M. (2015) Sequel to "The Life Histories of Asian Butterflies Vols. I & II". Yoshiko Igarashi, Tokyo, 355 pp.

    Iturralde-Vinent, M.A. & MacPhee, R.D.E. (1996) Age and paleographical origin of Dominican amber. Science, 273, 1850–1852.
    https://doi.org/10.1126/science.273.5283.1850

    Jarzembowski, E.A. (1980) Fossil insects from the Bembridge Marls, Palaeogene of the Isle of Wight, Southern England. Bulletin of the British Museum (Natural History), Geology, 33, 237–293.

    Jeratthitikul, E., Lewvanich, A., Butcher, B.A. & Lekprayoon, C. (2009) A taxonomic study of the genus Eurema Hübner, [1819] (Lepidoptera: Pieridae) in Thailand. Natural History Journal Chulalongkorn University, 9, 1–20.

    Kandul, N.P., Lukhtanov, V.A., Dantchenko, A.V., Coleman, J.A.S., Sekercioglu, C.H., Haig, D. & Pierce, N.E. (2004) Phylogeny of Agrodiaetus Hübner 1822 (Lepidoptera: Lycaenidae) inferred from mtDNA sequences of COI and COII and nuclear sequences of EF1-α: karyotype diversification and species radiation. Systematic Biology, 53, 278–298.
    https://doi.org/10.1080/10635150490423692

    Kawahara, A.Y. & Breinholt, J.W. (2014) Phylogenomics provides strong evidence for relationships of butterflies and moths. Proceedings of the Royal Society B, 281, 20140970.
    https://doi.org/10.1098/rspb.2014.0970

    Kawahara, A.Y. (2009) Phylogeny of snout butterflies (Lepidoptera: Nymphalidae: Libytheinae): combining evidence from the morphology of extant, fossil, and recently extinct taxa. Cladistics, 25, 263–278.
    https://doi.org/10.1111/j.1096-0031.2009.00251.x

    Kawahara, A.Y. (2013) Systematic revision and review of the extant and fossil snout butterflies (Lepidoptera: Nymphalidae: Libytheinae). Zootaxa, 3631 (1), 1–74.
    https://doi.org/10.11646/zootaxa.3631.1.1

    Kernbach, K. (1967) Über die bisher im Pliozän von Willershausen gefundenen Schmetterlings- und Raupenreste. Berichte Naturhistorisches Gesellschaft Hannover, 111, 103–108.

    Kirby, W.F. (1871) A Synonymic Catalogue of Diurnal Lepidoptera. John van Voorst, London, vii + 690 pp.

    Kirby, W.F. (1877) A Synonymic Catalogue of Diurnal Lepidoptera. Supplement. John van Voorst, London, 193 pp.
    https://doi.org/10.5962/bhl.title.11413

    Klots, A.B. (1931) A generic revision of the Pieridae (Lepidoptera). Entomologica Americana, 12, 139–204.

    Kozak, K.M., Wahlberg, N., Neild, A.F.E., Dasmahapatra, K.J.K. & Jiggins, C.D. (2015) Multilocus species trees show the recent adaptive radiation of the mimetic Heliconius butterflies. Systematic Biology, 64, 505–524.
    https://doi.org/10.1093/sysbio/syv007

    Kozlov, M.V. (1988) Paleontology of lepidopterans and problems of the phylogeny of the order Papilionida. In: Ponomarenko, A.G. (Ed.), The Mesozoic-Cenozoic Crisis in the Evolution of Insects. Academy of Sciences, Moscow, pp. 16–69. [in Russian]

    Kristensen, N.P. (1976) Remarks on the family-level phylogeny of butterflies (Insecta, Lepidoptera, Rhopalocera). Zeitschrift für zoologische Systematik und Evolutionsforschung, 14, 25–33.
    https://doi.org/10.1111/j.1439-0469.1976.tb00515.x

    Kristensen, N.P., Scoble, M.J. & Karsholt, O. (2007) Lepidoptera phylogeny and systematics: the state of inventorying moth and butterfly diversity. In: Zhang, Z.-Q. & Shear, W.A. (Eds.), Linnaeus Tercentenary: Progress in Invertebrate Taxonomy. Zootaxa, 1668, pp. 1–766.

    Kukalová-Peck, J. (1991) Fossil history and the evolution of Hexapod structures. In: CSIRO, Division of Entomology (Ed.), The insects of Australia. Vol. 1. 2nd Edition. CSIRO, Melbourne, pp. 141–179.

    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

    Labandeira, C.C. (2014) Amber. In: Laflamme, M., Schiffbauer, J.D. & Parroch, S.A. (Eds.), Reading and writing of the Fossil Record: Preservational Pathways to Exceptional Preservation. Paleontological Society Papers, 20, 163–216.

    Labandeira, C.C., Yang, Q., Santiago-Blay, J.A., Hotton, C.L., Monteiro, A., Wang, Y.-J., Goreva, Y., Shih, C.-K., Siljeström, S., Rose, T.R., Dilcher, L.D. & Ren, D. (2016) The evolutionary convergence of mid-Mesozoic lacewings and Cenozoic butterflies. Proceedings Royal Society B, 283, 1–9.
    https://doi.org/10.1098/rspb.2015.2893

    Larsen, T.B. (1974) Butterflies of Lebanon. National Council for Scientific Research, Beirut, xv + 255 pp.

    Larsen, T.B. (2005) Butterflies of West Africa. Text Volume. Apollo Books, Stenstrup, 595 pp.

    Larsson, S.G. (1978) Baltic Ambera Palaeobiological Study. Scandinavian Science Press, Klampenborg, 192 pp.

    Leestmans, R. (1983) Les Lépidoptères fossiles trouvés en France (Insecta Lepidoptera). Linneana belgica, 9, 64–89.

    Lutz, H. (1984) Beitrag zur Kenntnis der Unteroligozänen Insektenfauna von Ceresté [sic] (Süd-Frankreich). Documenta naturae, 21, 1–26.

    MacFadden, B.J. (1992) Fossil Horses. Systematics, Paleobiology, and Evolution of the Family Equidae. Cambridge University Press, Cambridge, xii + 369 pp.

    Magallón, S. (2014) A review of the effect of relaxed clock method, long branches, genes, and calibrations in the estimation of angiosperm age. Botanical Sciences, 92, 1–22.
    https://doi.org/10.17129/botsci.37

    Marshall, C.R. (1997) Confidence intervals on stratigraphic ranges with nonrandom distributions of fossils. Paleobiology, 23, 165–173.
    https://doi.org/10.1017/S0094837300016766

    Martínez-Delclòs, X., Briggs, D.E.G. & Peñalver, E. (2004) Taphonomy of insects in carbonates and amber. Palaeogeography, Palaeoclimatology, Palaeoecology, 203, 19–64.
    https://doi.org/10.1016/S0031-0182(03)00643-6

    Martins Neto, R.G. (1989) Novos insetos terciários do Estado de São Paulo. Revista Brasileira de Geociências, 19, 375–386.

    Martins Neto, R.G., Kucera-Santos, J.C., Vieira, F.R.de M. & Fragoso, L.M. de C. (1993) Nova espécie de borboleta (Lepidoptera: Nymphalidae: Satyrinae) da formação Tremembé, Oligoceno do Estado de São Paulo. Acta Geologica Leopoldensia, 37, 5–16.

    Megens, H.-J., van Moorsel, C.H.M., Piel, W.H., Pierce, N.E. & Jong, R. de (2004) Tempo of speciation in a butterfly genus from the Southeast Asian tropics, inferred from mitochondrial and nuclear DNA sequence data. Molecular Phylogenetics and Evolution, 31, 1181–1196.
    https://doi.org/10.1016/j.ympev.2003.10.002

    Miller, J.S. (1987) Phylogenetic studies in the Papilioninae (Lepidoptera: Papilionidae). Bulletin American Museum Natural History, 186, 365–512.

    Miller, J.Y. & Brown, F.M. (1989) A new Oligocene fossil butterfly, Vanessaamerindica (Lepidoptera: Nymphalidae), from the Florissant Formation, Colorado. Bulletin Allyn Museum, 126, 1–9.

    Miller, J.Y., Miller, L.D. & Ivie, M.A. (2012) A new lepidopteran fossil from the Canyon Ferry Reservoir deposit in southwestern Montana (Lepidoptera: Nymphalidae: Satyrinae). Bulletin Allyn Museum, 167, 1–12.

    Miller, L.D. (1968) The higher classification, phylogeny and zoogeography of the Satyridae (Lepidoptera). Memoirs American Entomological Society, 24, 1–174.
    https://doi.org/10.1130/MEM110-p1

    Miller, L.D. (1970) Nomenclature of wing veins and cells. Journal of Research on the Lepidoptera, 8 (1969), 37–48.

    Misof, B., Liu, S., Meusemann, K., Peters, R.S., Donath, A., Mayer, C., Frandsen, P.B., Ware, J., Flouri, T., Beutel, R.G., Niehuis, O., Petersen, M., Izquierdo-Carrasco, F., Wappler, T., Rust, J., Aberer, A.J., Aspöck, U., Aspöck, H., Bartel, D., Blanke, A., Berger, S., Böhm, A., Buckley, T.R., Calcott, B., Chen, J., Friedrich, F., Fukui, M., Fujita, M., Greve, C., Grobe, P., Gu, S., Huang, Y., Jermiin, L.S., Kawahara, A.Y., Krogmann, L., Kubiak, M., Lanfear, R., Letsch, H., Li, Y., Li, Z., Li, J., Lu, H., Machida, R., Mashimo, Y., Kapli, P., McKenna, D.D., Meng, G., Nakagaki, Y., Navarrete-Heredia, J.L., Ott, M., Ou, Y., Pass, G., Podsiadlowski, L., Pohl, H., von Reumont, B.M., Schütte, K., Sekiya, K., Shimizu, S., Slipinski, A., Stamatakis, A., Song, W., Su, X., Szucsich, N.U., Tan, M., Tan, X., Tang, M., Tang, J., Timelthaler, G., Tomizuka, S., Trautwein, M., Tong, X., Uchifune, T., Walzl, M.G., Wiegmann, B.M., Wilbrandt, J., Wipfler, B., Wong, T.K., Wu, Q., Wu, G., Xie, Y., Yang, S., Yang, Q., Yeates, D.K., Yoshizawa, K., Zhang, Q., Zhang, R., Zhang, W., Zhang, Y., Zhao, J., Zhou, C., Zhou, L., Ziesmann, T., Zou, S., Li, Y., Xu, X., Zhang, Y., Yang, H., Wang, J., Wang, J., Kjer, K.M. & Zhou, X. (2014) Phylogenomics resolves the timing and pattern of insect evolution. Science, 346, 763–767.
    https://doi.org/10.1126/science.1257570

    Munroe, E. (1961) The classification of the Papilionidae. Canadian Entomologist, 17 (Supplement), 1–51.

    Murata, Y. (1998) Notes on the evolutionary process of butterflies and butterfly fossils. Butterflies, 20 (part 1 & 2), 4–17 & 27–40. [in Japanese with English summary]

    Nazari, V., Zakharov, E.V. & Sperling, F.A.H. (2007) Phylogeny, historical biogeography, and taxonomic ranking of Parnassiinae (Lepidoptera: Papilionidae) based on morphology and seven genes. Molecular Phylogenetics and Evolution, 42, 131–156.
    https://doi.org/10.1016/j.ympev.2006.06.022

    Nekrutenko, Y.P. (1965) Tertiary nymphalid butterflies and some phylogenetic aspects of systematic lepidopterology. Journal of Research on the Lepidoptera, 4, 149–158.

    Nel, A. & Descimon, H. (1984) Une nouvelle espèce de Lépidoptère fossile du Stampien de Cereste (04) (Lepidoptera Satyridae). Géologie Méditerranéenne, 11, 287–293.

    Nel, A. & Nel, J. (1985) A propos de chenilles et de chrysalides fossiles du stampien en France, Alexanor, 14, 126–130.

    Nel, A. & Nel, J. (1986) A propos de deux Lépidoptères fossiles du Stampien d'Aix- en-Provence (Lepidoptera, Hesperiidae et Nymphalidae Satyrinae), Bulletin du Muséum National d' Histoire Naturelle, 8, 343–350.

    Nel, A., Nel, J. & Balme, C. (1993) Un nouveau Lépidoptère Satyrinae fossile de l’Oligocène du Sud-Est de la France (Insecta, Lepidoptera, Nymphalidae). Linneana belgica, 14, 20–36.

    Nielsen, E.S. & Common, I.F.B. (1991) Lepidoptera. In: CSIRO, Division of Entomology (Ed.), The insects of Australia. Vol. 2. 2nd Edition. CSIRO, Melbourne, pp. 817–915.

    Novacek, M.J. (1992) Fossils as critical data for phylogeny. In: Novacek, M.J. & Wheeler, Q.D. (Eds.), Extinction and Phylogeny. Columbia University Press, New York, pp. 46–88.

    O'Leary, M.A., Bloch, J.I., Flynn, J.J., Gaudin, T.J., Giallombardo, A., Giannini, N.P., Goldberg, S.L., Kraatz, B.P., Luo, Z-X., Meng, J., Ni, X., Novacek, M.J., Perini, F.A., Randall, Z.S., Rougier, G.W., Sargis, E.J., Silcox, M.T., Simmons, N.B., Spaulding, M., Velazco, P.M., Weksler, M.,Wible, J.R. & Cirranello, A.L. (2013) The Placental Mammal Ancestor and the Post–K-Pg Radiation of Placentals. Science, 339 (6120), 662–667.
    https://doi.org/10.1126/science.1229237

    Omoto, K., Katoh, T., Chichvarkhin, A. & Yagi, T. (2004) Molecular systematics and evolution of the "Apollo" butterflies of the genus Parnassius (Lepidoptera: Papilionidae), based on mitochondrial DNA sequence data. Gene, 326, 141–147.
    https://doi.org/10.1016/j.gene.2003.10.020

    Otaki, J.M., Kimura, Y. & Yamamoto, H. (2006) Molecular phylogeny and color-pattern evolution of Vanessa butterflies (Lepidoptera, Nymphalidae). Transactions of the Lepidopterological Society of Japan, 57, 359–370.

    Page, R.D.M. & Holmes, E.C. (1998) Molecular Evolution, a Phylogenetic Approach. Blackwell Science Ltd, Oxford, v + 346 pp.

    Paluch, M., Casagrande, M.M. & Mielke, O.H.H. (2003) Tampão genital de Actinote Hübner, como caráter toxonômico (Lepidoptera, Nymphalidae, Acraeinae). Revista Brasileira de Entomologia, 47, 573–580.
    https://doi.org/10.1590/S0085-56262003000400007

    Parham, J.F. Parham, J.F., Donoghue, P.C.J., Bell, C.J., Calway, T.D., Head, J.J., Holroyd, P.A., Inoue, J., Imris, R.B., Joyce, W.G., Ksepka, D.T., Patané, J.S.L., Smith, N.D., Tarver, J.E., van Tuinen, M., Yang, Z., Angielczyk, K.D., Greenwood, J.M., Hipsley, C.A., Jacobs, L., Makovicky, P.J., Müller, J., Smith, K.T., Theodor, J.M., Warnock, R.C.M. & Benton, M.J. (2012) Best practices for justifying fossil calibrations. Systematic Biology, 61, 346–359.

    Patterson, C. (1981) Significance of fossils in determining evolutionary relationships. Annual Revue Ecology Systematics, 12, 195–223.
    https://doi.org/10.1146/annurev.es.12.110181.001211

    Peña, C. & Wahlberg, N. (2008) Prehistorical climate change increased diversification of a group of butterflies. Biology Letters, 4, 274–278.
    https://doi.org/10.1098/rsbl.2008.0062

    Peñalver, E. & Grimaldi, D.A. (2006) New data on Miocene butterflies in Dominican Amber (Lerpidoptera: Riodinidae and Nymphalidae) with the description of a new nymphalid. American Museum Novitates, 3591, 1–17.
    https://doi.org/10.1206/0003-0082(2006)3519[1:NDOMBI]2.0.CO;2

    Penz, C.M., Freitas, A.V.L., Kaminski, L.A., Casagrande, M.M. & DeVries, P.J. (2013) Adult and early-stage characters of Brassolini contain conflicting phylogenetic signal (Lepioptera, Nymphalidae). Systematic Entomology, 38, 316–333.
    https://doi.org/10.1111/syen.12000

    Pfretzschner, H.U. (1998) Ein weitertes Exemplar von Lethe ? corbieri Nel 1993 (Lepidoptera, Satyridae) aus dem Unter-Oligozän von Céreste (Süd-Frankreich). Paläontologische Zeitschrift, 72, 59–64.
    https://doi.org/10.1111/syen.12000

    Poinar, G.O. (1992) Life in Amber. Stanford University Press, Stanford, xiii + 350 pp.

    Pongrácz, A. (1928) Die fossilen Insekten von Ungarn. Annales historico-naturalis Musei nationalis Hungarici, 25, 91–194.

    Ponomarenko, A.G. & Schultz, O. (1988) Kataloge der wissenschaftlichen Sammlung des Naturhistorischen Museums in Wien. Bd. 6. Paläozoologie. Heft 1. Natuurhistorisches Museum, Wien, 39 pp., 14 pls.

    Prothero, D.R. & Berggren, W.A. (Eds.) (1992) EoceneOligocene Climatic and Biotic Evolution. Princeton University Press, New Jersey, xiv + 568 pp.

    Prothero, D.R. (1994) The EoceneOligocene Transition. Paradise lost. Columbia University Press, New York, xviii + 291 pp.

    Rand, D.B., Heath, A., Suderman, T. & Pierce, N.E. (2000) Phylogeny and life history evolution of the genus Chrysoritis within the Aphnaeini (Lepidoptera: Lycaenidae), inferred from mitochondrial cytochrome oxidase I sequences. Molecular Phylogenetics and Evolution, 17, 85–96.
    https://doi.org/10.1006/mpev.2000.0820

    Rebel, H. (1898) Fossile Lepidopteren aus der Miocän-Formation von Gabbro. Sitzungsberichte der Akademie der Wissenschaften Wien, 107, 731–745.

    Regier, J. C. & 19 others (2009) Toward reconstructing the evolution of advanced moths and butterflies (Lepidoptera: Ditrysia): an initial molecular study. BMC Evolutionary Biology, 9, 280.
    https://doi.org/10.1186/1471-2148-9-280

    Rodriguez-Trelles, F., Tarrio, R. & Ayala, F.J. (2002) A methodological bias toward overestimation of molecular evolutionary time scales. Proceedings of the National Academy of Sciences, U.S.A., 99, 8112–8115.
    https://doi.org/10.1073/pnas.122231299

    Salemi, M. & Vandamme, A.-M. (Eds.), (2003) The Phylogenetic Handbook. A Practical Approach to DNA and Protein Phylogeny. Cambridge University Press, Cambridge, 406 pp.

    Schatz, E. & Röber, J. (1892) Die Familien und Gattungen der Tagfalter Systematisch und Analytisch Bearbeitet. Löwensohn, Fürth, ii + 284 pp.

    Schwanwitsch, B.N. (1943) Wing pattern in papilionid Lepidoptera. Entomologist, 76, 201–203.

    Scoble, M.J. (1992) The LepidopteraForm, Function and Diversity. Oxford University Press, Oxford, xi + 404 pp.

    Scott, J.A. (1986) The Butterflies of North America. Stanford University Press, Stanford, xiii + 583 pp.

    Scudder, S.H. (1872) Description d’un nouveau papillon fossile (Satyrites Reynesii) trouvé à Aix en Provence. Revue et Magasin de Zoologie, 23, (2), 66–72.cudder, S.H. (1875) Fossil butterflies. Memoirs American Association Advancement Science, 1, 1–99.

    Scudder, S.H. (1878) [Prodryas persephone]. Bulletin United States Geological Survey, 4, 524–526.

    Scudder, S.H. (1889) The Fossil butterflies of Florissant. United States Geological Survey, 8th Annual Report, 439–472.

    Scudder, S.H. (1892) Some insects of special interest from Florissant, Colorado, and other points in the Tertiaries of Colorado and Utah. Bulletin United States Geological Survey, 93, 1–35.

    Shields, O. (1976) Fossil butterflies and the evolution of Lepidoptera. Journal of Research on the Lepidoptera, 15, 132–143.

    Shields, O. (1985) Southeast Asian affinities in Colorado Oligocene Libytheidae. Tokurana (Acta Rhopalocera), Special Volume 1, 13–24.

    Silva-Brandão, K.L., Freitas, A.V.L., Brower, A.V.Z. & Solferini, V.N. (2005) Phylogenetic relationships of the New World Troidini swallowtails (Lepidoptera: Papilionidae) based on COI, COII, and EF-1a genes. Molecular Phylogenetics and Evolution, 36, 468–483.
    https://doi.org/10.1016/j.ympev.2005.04.007

    Silva-Brandão, K.L., Wahlberg, N., Francini, R.B., Azeredo-Espin, A.M.L., Brown Jr., K.S., Paluch, M., Lees, D.C. & Freitas, A.V.L. (2008) Phylogenetic relationships of butterflies of the tribe Acraeini (Lepidoptera, Nymphalidae, Heliconiinae) and the evolution of host plant use. Molecular Phylogenetics and Evolution, 46, 515–531.
    https://doi.org/10.1016/j.ympev.2007.11.024

    Simonsen, T.J., Zakharov, E.V., Djernaes, M., Cotton, A.M., Vane-Wright, R.I. & Sperling, F.A.H. (2011) Phylogenetics and divergence times of Papilionae (Lepidoptera) with special reference to the enigmatic genera Teinopalpus and Meandrusa. Cladistics, 27, 113–137.
    https://doi.org/10.1111/j.1096-0031.2010.00326.x

    Skalski, A.W. (1976) Les lépidoptères fossiles de l’ambre. Etat actuel de nos connaissances (2e partie). Linneana belgica, 6, 195–208.

    Smith, A.G., Smith, D.G. & Funnell, B.M. (1994) Atlas of Mesozoic and Cenozoic Coastlines. Cambridge University Press, Cambridge, ix + 99.

    Smith, M.E., Carroll, A.R., & Singer, B.S. (2008). Synoptic reconstruction of a major lake system: Eocene Green River Formation, Western United States. Geological Society of America Bulletin, 120, 54–84.
    https://doi.org/10.1130/B26073.1

    Sober, E. (1988) Reconstructing the Past. MIT Press, Cambridge (Mass.), London, xiii + 265.

    Sohn, J.-C., Labandeira, C., Davis, D. & Mitter, C. (2012) An annotated catalog of fossil and subfossil Lepidoptera (Insecta: Holometabola) of the world. Zootaxa, 3286, 1–132.

    Stichel, H. (1907) Lepidoptera Rhopalocera, Fam. Papilionidae, Subfam. Parnassiinae. Genera Insectorum, 58, 1–60.

    Talavera, G., Lukhtanov, V.A., Pierce, N.E. & Vila, R. (2013) Establishing criteria for higher-level classification using molecular data: the systematics of Polyommatus blue butterflies (Lepidoptera: Lycaenidae). Cladistics, 29, 166–192.
    https://doi.org/10.1111/j.1096-0031.2012.00421.x

    Théobald, N. (1937) Note complémentaires sur les insectes fossiles oligocènes des gypses d’Aix-en-Provence. Bulletin de la Société des Sciences de Nancy, 6, 157–179.

    Timmermans, M.J.T.N., Lees, D.C. & Simonsen, T.J. (2014) Towards a mitogenomic phylogeny of Lepidoptera. Molecular Phylogenetics and Evolution, 79, 169–178.
    https://doi.org/10.1016/j.ympev.2014.05.031

    Tree of Life Web Project (2010). Ditrysia. Version 17 November 2010. Available from: http://tolweb.org/Ditrysia/11868/2010.11.17 (accessed 23 May 2015)

    Trewick, S.A. (2000) Molecular evidence for dispersal rather than vicariance as the origin of flightless insect species on the Chatham Islands, New Zealand. Journal of Biogeography, 27, 1189–1200.
    https://doi.org/10.1046/j.1365-2699.2000.00492.x

    Turati, E. (1918) Variabilità del Parnassius apollo pumilus Stich. a ricerche sull' origine dei Parnassius. Atti Società Italiana di Scienza naturali, 57, 29–89.

    Ugelvig, L.V., Vila, R., Pierce, N.E. & Nash, D.R. (2011) A phylogenetic revision of the Glaucopsyche section (Lepidoptera: Lycaenidae), with special focus on the Phengaris-Maculinea clade. Molecular Phylogenetics and Evolution, 61, 237–243.
    https://doi.org/10.1016/j.ympev.2011.05.016

    Urbahn, E. & Urbahn, H. (1939) Die Schmetterlinge Pommerns mit einem vergleichenden Überblick über den Ostseeraum. Stettiner entomologische Zeitung, 100–101, 185–826.

    van Schepdael, J. (1974) Macrolépidoptères fossiles du domain paléarctique—la paléontologie au service de l’Écologie. Naturalistes belges, 55, 3–37.

    Vane-Wright, R.I. & Hughes, H.W.D. (2007) Did a member of the Vanessa indica complex (Nymphalidae) formerly occur in North America? Journal of the Lepidopterists' Society, 61, 199–212.

    Vane-Wright, R.I. (2004) Butterflies at that awkward age. Nature, 428, 477–480.
    https://doi.org/10.1038/428477a

    Verity, R. (1947) Le farfalle diurne d’Italia. Vol. 3. Marzocco, Firenze, xvi + 318 pp.

    Verity, R. (1952) Les variations géographiques et saisonnières des papillons diurnes en France. Vol. 2. Revue française de Lépidopterologie, 1952 (Supplément), 165 pp.

    von Heyden, C. (1859) Fossile Insekten aus der Rheinischen Braunkohle. Palaeontographica, 8, 1–15.

    Wahlberg, N. (2006) That awkward age for butterflies: insights from the age of the butterfly subfamily Nymphalinae. Systematic Biology, 55, 703–714.
    https://doi.org/10.1080/10635150600913235

    Wahlberg, N., Braby, M.F., Brower, A.V.Z., de Jong, R., Lee, M.-M., Nylin, S., Pierce, N.E., Sperling, F.A.H., Vila, R.U., Warren, A.D. & Zakharov, E. (2005a) Synergetic effects of combining morphological and molecular data in resolving the phylogeny of butterflies and skippers. Proceedings of the Royal Society B, 272, 1577–1586.
    https://doi.org/10.1098/rspb.2005.3124

    Wahlberg, N. & Brower, A.V.Z. (2006) Satyrinae Boisduval 1833. Version 09 November 2006 (under construction). Available from: http://tolweb.org/Satyrinae/12189/2006.11.09 in The Tree of Life Web Project. Available from: http://tolweb.org/ (accessed 23 May 2015)

    Wahlberg, N., Brower, A.V.Z. & Nylin, S. (2005b) Phylogenetic relationships and historical biogeography of tribes and genera in the subfamily Nymphalinae (Lepidoptera: Nymphalidae). Biological Journal Linnean Society, 86, 227–251.
    https://doi.org/10.1111/j.1095-8312.2005.00531.x

    Wahlberg, N. & Freitas, A.V.L. (2007) Colonization of and radiation in South America by butterflies in the subtribe Phyciodina (Lepidoptera: Nymphalidae). Molecular Phylogenetics and Evolution, 44, 1257–1272.
    https://doi.org/10.1016/j.ympev.2007.04.012

    Wahlberg, N., Leneveu, J., Kodandaramaiah, U., Peña, C., Nylin, S., Freitas, A.V.L. & Brower, A.V.Z. (2009) Nymphalid butterflies diversify following near demise at the Cretaceous/Tertiary boundary. Proceedings Royal Society B, 276, 4295–4302.
    https://doi.org/10.1098/rspb.2009.1303

    Wahlberg, N. & Peña, C. (2015) Systematics and Evolution of Nymphalidae. Available from: http://www.nymphalidae.net/(accessed 12 November 2015)

    Wahlberg, N., Rota, J., Braby, M.F. & Pierce, N. (2014) Revised systematics and higher classification of pierid butterflies (Lepidoptera: Pieridae) based on molecular data. Zoologica Scripta, 43, 641–650.
    https://doi.org/10.1111/zsc.12075

    Walker, J.D., Geissman, J.W., Bowring, S.A. & Babcock, L.E. (2013) The Geological Society of America Geologic Time Scale. Bulletin of the Geological Society of America, 125, 259–272.
    https://doi.org/10.1130/B30712.1

    Wangrin, G. (1939) Tagfalter-Versteinerung in einer “Stettiner Kugel”. Entomologische Zeitschrift, 53, 192–194.

    Warren, A.D., Ogawa, J.R. & Brower, A.V.Z. (2008) Phylogenetic relationships of subfamilies and circumscription of tribes in the family Hesperiidae (Lepidoptera: Hesperioidea). Cladistics, 24, 642–676.
    https://doi.org/10.1111/j.1096-0031.2008.00218.x

    Warren, A.D., Ogawa, J.R. & Brower, A.V.Z. (2009) Revised classification of the family Hesperiidae (Lepidoptera: Hesperioidea) based on combined molecular and morphological data. Systematic Entomologist, 34, 467–523.
    https://doi.org/10.1111/j.1365-3113.2008.00463.x

    Westwood, J.O. (1854) Contributions to Fossil Entomology. Quarterly Journal of the Geological Society of London, 10, 378–396.
    https://doi.org/10.1144/GSL.JGS.1854.010.01-02.43

    Wiemers, M., Stradomsky, B.V. & Vovolazhsky, D.L. (2010) A molecular phylogeny of Polyommatus s.str. and Plebicula based on mitochondrial COI and nuclear ITS2 sequences (Lepidoptera: Lycaenidae). European Journal of Entomology, 107, 325–336.
    https://doi.org/10.14411/eje.2010.041

    Wiley, E.O. (1981) Phylogenetics. The theory and practice of phylogenetic systematics. John Wiley & Sons, New York, etc., xv + 439 pp.

    Wilf, P. & Escapa, H. (2014) Green Web or megabiased clock? Plant fossils from Gondwanan Patagonia speak on evolutionary radiations. New Phytologist, 207, 283–290.
    https://doi.org/10.1111/nph.13114

    Willmott, K.R. & Freitas, A.V.L. (2006) Higher-level phylogeny of the Ithomiinae (Lepidoptera: Nymphalidae): classification, patterns of larval hostplant colonization and diversification. Cladistics, 22, 297–368.
    https://doi.org/10.1111/j.1096-0031.2006.00108.x

    Woodruff, R.E. (1994) Life or death in amber? Insecta Mundi, 8, 137–142.

    Wootton, R.J. (1979) Function, homology and terminology in insect wings. Systematic Entomology, 4, 81–93.
    https://doi.org/10.1111/j.1365-3113.1979.tb00614.x

    Zakharov, E.V., Caterino, M.S. & Sperling, F.A. (2004) Molecular Phylogeny, Historical Biogeography, and Divergence Time Estimates for Swallowtail Butterflies of the Genus Papilio (Lepidoptera: Papilionidae). Systematic Biology, 53, 1–23.
    https://doi.org/10.1080/10635150490423403

    Zeuner, F.E. (1930) Eine neue tertiäre Insektenfauna aus Süddeutschland. Natur und Museum, Frankfurt, 60, 270–281.

    Zeuner, F.E. (1942) Two new fossil butterflies of the family Pieridae. Annals and Magazine of Natural History, Series 11, 9, 409–416.
    https://doi.org/10.1080/03745481.1942.9755493

    Zeuner, F.E. (1961) Notes on the evolution of the Rhopalocera (Lep.). Verhandlungen des XI. Internationalen Kongresses für Entomologie, 1 (1960), 310–313.