Nymphalidae Systematics Group

NSG publications

Publications by members of the NSG. For full publication lists of individual researchers see People-pages

2012

  1. Condamine, F. L., Sperling, F. A. H., Wahlberg, N., Rasplus, J.-Y. & Kergoat, G. J. 2012: What causes latitudinal gradients in species diversity? Evolutionary processes and ecological constraints on swallowtail biodiversity. Ecology Letters 15: 267-277. doi:10.1111/j.1461-0248.2011.01737.x
  2. Heikkilä, M., Kaila, L., Mutanen, M., Peña, C. & Wahlberg, N. 2012: Cretaceous origin and repeated Tertiary diversification of the redefined butterflies. Proceedings of the Royal Society Series B Biological Sciences 279: 1093-1099. doi:10.1098/rspb.2011.1430
  3. Kodandaramaiah, U., Konvicka, M., Tammaru, T., Wahlberg, N. & Gotthard, K. 2012: Phylogeography of the woodland brown butterfly Lopinga achine (Nymphalidae: Satyrinae): implications for conservation. Journal of Insect Conservation, in press.
  4. Núñez Aguila, R., Oliva Plasencia, E., Matos Maravi, P. F. & Wahlberg, N. 2012: Cuban Calisto (Lepidoptera: Nymphalidae: Satyrinae), a review based on morphological and DNA data. ZooKeys 165: 57-105. doi:10.3897/zookeys.165.2206
  5. Zahiri, R., Holloway, J.D., Kitching, I.J., Lafontaine, J.D., Mutanen, M. & Wahlberg, N. 2012: Molecular phylogenetics of Erebidae (Lepidoptera, Noctuoidea). Systematic Entomology 37: 102-124. doi:10.1111/j.1365-3113.2011.00607.x

2011

  1. de Jong, M. A., Wahlberg, N., van Eijk, M., Brakefield, P.M. & Zwaan, B.J. 2011: Limited mitochondrial DNA differentiation for range-wide populations of Bicyclus anynana suggests a rapid expansion from recent refugia. PLoS ONE 6: e21385. doi:10.1371/journal.pone.0021385
  2. Kodandaramaiah, U. 2011: Tectonic calibrations in molecular dating. Current Zoology 57: 116-124. pdf
  3. Marin, M. A., Peña, C., Freitas, A. V. L., Wahlberg, N. & Uribe, S. I. 2011: From the phylogeny of the Satyrinae butterflies to the systematics of Euptychiina (Lepidoptera, Nymphalidae): history, progress and prospects. Neotropical Entomology 40: 1-13.
  4. Mullen, S. P., Savage, W. K., Wahlberg, N. & Willmott, K. 2011: Rapid diversification and not clade age explains high diversity in neotropical Adelpha butterflies. Proceedings of the Royal Society Series B Biological Sciences 278: 1777-1785. doi:10.1098/rspb.2010.2140
  5. Peña, C. 2011: Métodos de inferencia filogenética. Revista peruana de Biología 18: 265-267. [Commentary] pdf
  6. Peña, C., Nylin, S. & Wahlberg, N. 2011: The radiation of Satyrini butterflies (Nymphalidae: Satyrinae): a challenge for phylogenetic methods. Zoological Journal of the Linnean Society 161: 64-87. doi:10.1111/j.1096-3642.2009.00627.x
  7. Penz, C. M., Mohammadi, N. & Wahlberg, N. 2011: The Neotropical butterfly genus Blepolenis: wing pattern elements, phylogeny, and Pleistocene diversification (Lepidoptera, Nymphalidae). Zootaxa 2897: 1-17.
  8. Pulido-B., H. W., Andrade-C., M. G., Peña, C., & Lamas, G. 2011: Two new taxa of Euptychia Hübner, 1818 (Lepidoptera: Nymphalidae: Satyrinae) from the Andes of Colombia and Peru. Zootaxa 2906: 43-51.
  9. van Nieukerken, E.J., Kaila, L., Kitching, I.J., Kristensen, N.P., Lees, D.C., Minet, J., Mitter, C., Mutanen, M., Regier, J.C., Simonsen, T.J., Wahlberg, N., Yen, S.-H., Zahiri, R., Adamski, D., Baixeras, J., Bartsch, D., Bengtsson, B.Å., Brown, J.W., Bucheli, S.R., Davis, D.R., De Prins, J., De Prins, W., Epstein, M.E., Gentili-Poole, P., Gielis, C., Hättenschwiler, P., Hausmann, A., Holloway, J.D., Kallies, A., Karsholt, O., Kawahara, A., Koster, J.C., Kozlov, M. V., Lafontaine, J. D., Lamas, G., Landry, J.-F., Lee, S., Nuss, M., Park, K.-T., Penz, C., Rota, J., Schmidt, B. C., Schintlmeister, A., Sohn, J. C., Solis, M. A., Tarmann, G. M., Warren, A. D., Weller, S., Yakovlev, V. R., Zolotuhin, V. V. & Zwick, A. 2011: Order Lepidoptera. In: Zhang, Z.-Q. (Ed.), Animal biodiversity: An outline of higher-level classification and survey of taxonomic richness. Zootaxa 3148: 212-221. pdf.
  10. Veijalainen, A., Broad, G. R., Wahlberg, N., Longino, J. T., Sääksjärvi, I. E. 2011: DNA barcoding and morphology reveal two common species in one: Pimpla molesta stat. rev. separated from P. croceipes (Hymenoptera: Ichneumonidae). ZooKeys 124: 59-70. doi:10.3897/zookeys.124.1780
  11. Wahlberg, N. & Rubinoff, D. 2011: Vagility across Vanessa (Lepidoptera: Nymphalidae): mobility in butterfly species does not inhibit the formation and persistence of isolated sister taxa. Systematic Entomology 36: 362-370. doi:10.1111/j.1365-3113.2010.00566.x
  12. Wahlberg, N. & Rubinoff, D. 2011: Anomalous areas and awkward ages: alleviating concerns. Systematic Entomology 36: 604-606. doi:10.1111/j.1365-3113.2011.00590.x
  13. Xia, H., Wahlberg, N., Belák, S., Meng, X. J. & Liu, L. 2011: The emergence of genotypes 3 and 4 hepatitis E virus in swine and human: a phylogenetic perspective. Archives of Virology 156: 121-124. doi:10.1007/s00705-010-0818-6
  14. Xia, H., Wahlberg, N., Qiu, H.-J., Belák, S., Widén, F. & Liu, L. 2011: No phylogenetic evidence to support Shimen strain as the parental strain of the lapinized Chinese strain (C-strain) vaccine against classical swine fever. Archives of Virology 156: 1041-1044. doi:10.1007/s00705-011-0948-5
  15. Zahiri, R., Kitching, I. J., Lafontaine, J. D., Mutanen, M., Kaila, L., Holloway, J. D. & Wahlberg, N. 2011: A new molecular phylogeny offers hope for a stable family-level classification of the Noctuoidea (Lepidoptera). Zoologica Scripta 40: 158-173. doi:10.1111/j.1463-6409.2010.00459.x

2010

  1. 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. doi:10.1080/14772001003626814 pdf
  2. Kodandaramaiah, U. 2010: Use of Dispersal-Vicariance analysis in biogeography - a critique. Journal of Biogeography 37: 3-11. doi:10.1111/j.1365-2699.2009.02221.x
  3. Kodandaramaiah, U., Lees, D. C., Müller, C. J., Torres, E., Karanth, P. K. & Wahlberg, N. 2010: Phylogenetics and biogeography of a spectacular Old World radiation of butterflies: the subtribe Mycalesina (Lepidoptera: Nymphalidae: Satyrini). BMC Evolutionary Biology 10: 172. doi:10.1186/1471-2148-10-172 pdf
  4. Kodandaramaiah, U., Peña, C., Braby, M., Grund, R., Müller, C. J., Nylin, S. & Wahlberg, N. 2010: Phylogenetics of Coenonymphina (Nymphalidae: Satyrinae) and the problem of rooting rapid radiations. Molecular Phylogenetics and Evolution 54: 386-394. doi:10.1016/j.ympev.2009.08.012 pdf
  5. Liu, L., Xia, H., Baule, C., Belák, S. & Wahlberg, N. 2010: Effects of methodology and analysis strategy on robustness of pestivirus phylogeny. Virus Research 147: 47-52. doi:10.1016/j.virusres.2009.10.004 pdf
  6. Müller, C. J., Wahlberg, N. & Beheregaray , L. B. 2010: 'After Africa' - The evolutionary history and systematics of the genus Charaxes Ochsenheimer (Lepidoptera: Nymphalidae) in the Indo-Pacific Region. Biological Journal of the Linnean Society 100: 457-481. doi:10.1111/j.1095-8312.2010.01426.x pdf
  7. Mutanen, M., Wahlberg, N. & Kaila, L. 2010: Comprehensive gene and taxon coverage elucidates radiation patterns in moths and butterflies. Proceedings of the Royal Society Series B Biological Sciences 277: 2839-2848. doi:10.1098/rspb.2010.0392 pdf
  8. Peña, C., Nylin, S., Freitas, A. V. L. & Wahlberg, N. 2010: Biogeographic history of the butterfly subtribe Euptychiina (Lepidoptera, Nymphalidae, Satyrinae). Zoologica Scripta 39: 243-258. doi:10.1111/j.1463-6409.2010.00421.x pdf
  9. Salokannel, J., Rantala, M., & Wahlberg, N. 2010: DNA-barcoding clarifies species definitions of Finnish Apatania (Trichoptera: Apataniidae). Entomologica Fennica 21: 1-11. pdf
  10. Simonsen, T. J., Wahlberg, N., Warren, A. D. & Sperling, F. A. H. 2010: The evolutionary history of Boloria (Lepidoptera: Nymphalidae): phylogeny, zoogeography and larval foodplant relationships. Systematics and Biodiversity 8: 513-529. doi:10.1080/14772000.2010.532833 pdf
  11. Wahlberg, N., Snäll, N., Viidalepp, J., Ruohomäki, K. & Tammaru, T. 2010: The evolution of female flightlessness among Ennominae of the Holarctic forest zone (Lepidoptera, Geometridae). Molecular Phylogenetics and Evolution 55: 929-938. doi:10.1016/j.ympev.2010.01.025 pdf

2009

  1. Aduse-Poku, K., Vingerhoedt, E. & Wahlberg, N. 2009: Out-of-Africa again: a phylogenetic hypothesis of the genus Charaxes (Lepidoptera: Nymphalidae) based on 5 gene regions. Molecular Phylogenetics and Evolution 53: 463-478. doi:10.1016/j.ympev.2009.06.021 pdf
  2. Kodandaramaiah, U. 2009: Eyespot evolution - phylogenetic insights from Junonia and related butterfly genera (Nymphalidae: Junoniini). Evolution & Development 11: 489-497. doi:10.1111/j.1525-142X.2009.00357.x.
  3. Kodandaramaiah, U. 2009: Vagility - the neglected component in historical biogeography. Evolutionary Biology 36: 327-335. doi:10.1007/s11692-009-9068-5 pdf.
  4. Kodandaramaiah, U., Vallin, A. & Wiklund, C. 2009: Fixed eyespot display in a butterfly thwarts attacking birds. Animal Behaviour 77: 1415-1419 doi:10.1016/j.anbehav.2009.02.018 pdf.
  5. Kodandaramaiah, U. & Wahlberg, N. 2009: Phylogeny and biogeography of Coenonympha butterflies (Nymphalidae: Satyrinae) - patterns of colonization in the Holarctic. Systematic Entomology 34:315-323 doi:10.1111/j.1365-3113.2008.00453.x pdf.
  6. Lehmann, L., D. Stadie & R. Zahiri. 2009: Zum Winteraspekt der Makrolepidopterenfauna Südiran mit Anmerkungen zur Biologie einiger Arten (Lepidoptera: Bombycoidea, Papilionoidea, Geometroidea, Noctuoidea). Nachrichten des Entomologischen Vereins Apollo 30: 105-119.
  7. Leneveu, J., Chichvarkhin, A. & Wahlberg, N. 2009: Varying rates of diversification in the genus Melitaea (Lepidoptera: Nymphalidae) during the past 20 million years. Biological Journal of the Linnean Society 97:346-361. doi:10.1111/j.1095-8312.2009.01208.x pdf.
  8. Liu, L., Xia, H., Wahlberg, N., Belák, S. & Baule, C. 2009: Phylogeny, classification and evolutionary insights into pestiviruses. Virology 385:351-357. doi:10.1016/j.virol.2008.12.004 pdf.
  9. 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 of the Royal Society Series B Biological Sciences 276: 4295-4302. doi:10.1098/rspb.2009.1303 pdf Online supporting material can be downloaded here. Read ScienceNOW interview.
  10. Wahlberg, N., Weingartner, E., Warren, A. D. & Nylin, S. 2009: Timing major conflict between mitochondrial and nuclear genes in species relationships of Polygonia butterflies (Nymphalidae: Nymphalini). BMC Evolutionary Biology 9:92. doi:10.1186/1471-2148-9-92 pdf

2008

  1. Fibiger, M., G. Stangelmaier, C. Wieser & R. Zahiri 2008: Dichonia jahannamah, a new species from Iran in the subgenus Griposia Tams, 1939 (Lepidoptera, Noctuidae, Xyleninae). Nachrichten des Entomologischen Vereins Apollo 28: 121-124.
  2. Friberg, M., Bergman, M., Kullberg, J., Wahlberg, N. & Wiklund, C. 2008: Niche separation in space and time between two sympatric sister species - a case of ecological pleiotropy. Evolutionary Ecology 22: 1-18 pdf.
  3. Khaghaninia, S., S. A. Mohamadi, A. Sarafrazi, K. Hadade Iraninejad, E. Ebrahimi, S. Alavikia & R. Zahiri 2008: Geometric morphometric approach on the sexual dimorphism of Cydia pomonella (Lep.: Tortricidae) in the North West of Iran. Journal of Entomological Society of Iran 28: 51-62.
  4. Matov, A., R. Zahiri & J. D. Holloway 2008: The Heliothinae of Iran (Lepidoptera: Noctuidae). Zootaxa 1763: 1-37.
  5. Müller, G. C., V. D. Kravchenko, T. J. Witt, A. Junnila, J. Mooser, A. Saldaitis, K. Reshöft, P. Ivinskis, R. Zahiri & W. Speidel 2008: New Underwing taxa of the group of Catocala lesbia Christoph, 1887 (Lepidoptera, Noctuidae). Acta zoologica Lituanica 18: 30-49.
  6. Nylin, S. & Wahlberg, N. 2008: Does plasticity drive speciation? Host plant shifts and diversification in nymphaline butterflies (Lepidoptera: Nymphalidae) during the tertiary. Biological Journal of the Linnean Society 94: 115-130. doi:10.1111/j.1095-8312.2008.00964.x pdf.
  7. Peña, C., & N. Wahlberg 2008: Prehistorical climate change increased diversification of a group of butterflies. Biology Letters 4: 274-278. doi:10.1098/rsbl.2008.0062 pdf. Supplementary material. See commentary in Telegraph
  8. Silva Brandão, K. L., N. Wahlberg, R. B. Francini, A. M. L. Azeredo-Espin, K. S. Brown Jr., M. Paluch, D. C. Lees, and A. V. L. Freitas. 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 doi:10.1016/j.ympev.2007.11.024 pdf.
  9. Wahlberg, N., & Wheat, C. W. 2008: Genomic outposts serve the phylogenomic pioneers: designing novel nuclear markers for genomic DNA extractions of Lepidoptera. Systematic Biology 57: 231-242. doi:10.1080/10635150802033006 pdf, for list of primers see Molecular Methods or download Electronic Appendices.
  10. Zahiri, R. & M. Fibiger 2008: The Plusiinae of Iran (Lepidoptera: Noctuidae). SHILAP Revista de lepidopterología 36: 301-339.
  11. Zolotuhin, V. & R. Zahiri 2008: The Lasiocampidae of Iran (Lepidoptera). Zootaxa 1791: 1-52.

2007

  1. Brooks, D.R., Bilewitch, J., Condy, C., Evans, D.C., Folinsbee, K.E., Fröbisch, J., Halas, D., Hill, S., McLennan, D.A., Mattern, M., Tsuji, L.A., Ward, J.L., Wahlberg, N., Zamparo, D., Zanatta, D., 2007: Quantitative phylogenetic analysis in the 21st century. Revista Mexicana de Biodiversidad 78: 225-252 pdf.
  2. Brown Jr., K.S., Freitas, A.V.L., Wahlberg, N., von Schoultz, B., Saura, A.O., Saura, A. 2007: Chromosomal evolution in the South American Nymphalidae. Hereditas 144: 137-148. pdf
  3. 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. pdf.
  4. Gatesy, J., DeSalle, R. & Wahlberg, N. 2007: How many genes should a systematist sample? Conflicting insights from a phylogenomic matrix. Systematic Biology 56: 355-363 pdf.
  5. Kodandaramaiah, U. & Wahlberg, N. 2007: Out-of-Africa origin and dispersal mediated diversification of the butterfly genus Junonia (Nymphalidae: Nymphalinae). Journal of Evolutionary Biology 20: 2181-2191 pdf.
  6. Snäll, N., Tammaru, T., Wahlberg, N., Viidalepp, J., Ruohomäki, K., Savontaus, M.-L. & Huoponen, K. 2007: Phylogenetic relationships of the tribe Operophterini (Lepidoptera, Geometridae): a case study of the evolution of female flightlessness. Biological Journal of the Linnean Society 92: 241-252 pdf.
  7. Viidalepp, J., Tammaru, T., Snäll, N., Ruohomäki, K. & Wahlberg, N. 2007: Cleorodes Warren, 1894 does not belong in the tribe Boarmiini (Lepidoptera: Geometridae). European Journal of Entomology 104:303-309 pdf.
  8. 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 pdf.
  9. Wahlberg, N. & Saccheri, I. 2007: The effects of Pleistocene glaciations on the phylogeography of Melitaea cinxia (Lepidoptera: Nymphalidae). European Journal of Entomology 104: 675-684 pdf.
  10. Xia, H., Liu, L., Wahlberg, N., Baule, C. and Belák, S. 2007: Molecular phylogenetic analysis of bovine viral diarrhoea virus: a Bayesian approach. Virus Research 130:53-62 pdf.

2006

  1. Freitas, A. V. L. & C. Peña. 2006: Description of Genus Guaianaza for "Euptychia" pronophila (Lepidoptera: Nymphalidae: Satyrinae) with a description of the immature stages. Zootaxa 1163:49-59. pdf
  2. Janz, N., S. Nylin, & N. Wahlberg. 2006: Diversity begets diversity: host expansions and the diversification of plant-feeding insects. BMC Evolutionary Biology 6:4. pdf
  3. Peña, C., N. Wahlberg, E. Weingartner, U. Kodandaramaiah, S. Nylin, A. V. L. Freitas, and A. V. Z. Brower. 2006: Higher level phylogeny of Satyrinae butterflies (Lepidoptera: Nymphalidae) based on DNA sequence data. Molecular Phylogenetics and Evolution 40:29-49 pdf.
  4. Simonsen, T. J., Wahlberg, N., Brower, A. V. Z. & de Jong, R. 2006: Morphology, molecules and Fritillaries: approaching a stable phylogeny for Argynnini (Lepidoptera: Nymphalidae). Insect Systematics and Evolution 37:405-418 pdf.
  5. Wahlberg, N. 2006: That awkward age for butterflies: insights from the age of the butterfly subfamily Nymphalinae. Systematic Biology 55:703-714. pdf
  6. Weingartner, E., N. Wahlberg, & S. Nylin. 2006: Dynamics of host plant use and species diversity: a phylogenetic investigation in Polygonia butterflies (Nymphalidae). Journal of Evolutionary Biology 19:483-491 pdf.
  7. Weingartner, E., N. Wahlberg, & S. Nylin. 2006: Speciation in Pararge (Satyrinae: Nymphalidae) butterflies - North Africa is the source of ancestral populations of all Pararge species. Systematic Entomology 31:621-632 pdf.

2005

  1. Päivinen, J., Grapputo, A., Kaitala, V., Komonen, A., Kotiaho, J. S., Saarinen, K. & Wahlberg, N. 2005: Negative density-distribution relationship in butterflies. BMC Biology 3:5 pdf.
  2. Peña, C. & G. Lamas. 2005: Revision of the butterfly genus Forsterinaria Gray, 1973 (Lepidoptera: Nymphalidae, Satyrinae). Revista peruana biologia 12:5-48. pdf
  3. Wahlberg, N., M. F. Braby, A. V. Z. Brower, R. de Jong, M.-M. Lee, S. Nylin, N. Pierce, F. A. Sperling, R. Vila, A. D. Warren, & E. Zakharov. 2005: Synergistic effects of combining morphological and molecular data in resolving the phylogeny of butterflies and skippers. Proceedings of the Royal Society of London Series B Biological Sciences 272:1577-1586.pdf.
  4. Wahlberg, N., Brower, A. V. Z. & Nylin, S. 2005: Phylogenetic relationships and historical biogeography of tribes and genera in the subfamily Nymphalinae (Lepidoptera: Nymphalidae). Biological Journal of the Linnean Society 86:227-251. pdf.

2003

  1. Wahlberg, N., & Nylin, S. 2003: Morphology versus molecules: resolution of the positions of Nymphalis, Polygonia and related genera (Lepidoptera: Nymphalidae). Cladistics 19:213-223 pdf.
  2. Wahlberg, N., Oliveira, R. & Scott, J. A. 2003: Phylogenetic relationships of Phyciodes butterfly species (Lepidoptera: Nymphalidae): complex mtDNA variation and species delimitations. Systematic Entomology 28:257-273 pdf.
  3. Wahlberg, N., Weingartner, E. & Nylin, S. 2003: Towards a better understanding of the higher systematics of Nymphalidae (Lepidoptera: Papilionoidea). Molecular Phylogenetics and Evolution 28:473-484 pdf.

2002

  1. Wahlberg, N., Klemetti, T. & Hanski, I. 2002: Dynamic populations in a dynamic landscape: the metapopulation structure of the marsh fritillary butterfly. Ecography 25:224-232 pdf.
  2. Wahlberg, N., Klemetti, T., Selonen, V. & Hanski, I. 2002: Metapopulation structure and movements in five species of checkerspot butterflies. Oecologia 130:33-43 pdf.

2001

  1. Haikola, S., Fortelius, W., O'Hara, R., Kuussaari, M., Wahlberg, N., Saccheri, I. J., Singer, M. C. & Hanski, I. 2001: Inbreeding depression and the maintenance of genetic load in Melitaea cinxia metapopulations. Conservation Genetics 2:325-335 pdf.
  2. Wahlberg, N. 2001: On the status of the scarce fritillary Euphydryas maturna (Lepidoptera: Nymphalidae) in Finland. Entomologica Fennica 12:244-250 pdf.
  3. Wahlberg, N. 2001: The phylogenetics and biochemistry of host plant specialization in melitaeine butterflies (Lepidoptera: Nymphalidae). Evolution 55:522-537 pdf.
  4. Wahlberg, N., Kullberg, J. & Hanski, I. 2001: Natural history of some Siberian melitaeine butterfly species (Nymphalidae: Melitaeini) and their parasitoids. Entomologica Fennica 12:72-77 pdf.

2000

  1. Wahlberg, N. 2000: Comparative descriptions of the immature stages and ecology of five Finnish melitaeine butterfly species (Lepidoptera: Nymphalidae). Entomologica Fennica 11:167-174 pdf.
  2. Wahlberg, N. & Zimmermann, M. 2000: Pattern of phylogenetic relationships among members of the tribe Melitaeini (Lepidoptera: Nymphalidae) inferred from mtDNA sequences. Cladistics 16:347-363 pdf.
  3. Zimmermann, M., Wahlberg, N. & Descimon, H. 2000: Phylogeny of Euphydryas checkerspot butterflies (Lepidoptera: Nymphalidae) based on mitochondrial DNA sequence data. Annals of the Entomological Society of America 93:347-355 pdf.