Niklas Wahlberg

The circumscription of the subfamily Nymphalinae has changed many times over the past decades from encompassing about half of the ca. 6000 species of Nymphalidae to its current rather restricted sense of about 473 species (according to the latest version of the Classification of Nymphalidae). The species are placed into 6 tribes and 56 genera. The current tribes are Coeini, Nymphalini, Junoniini, Victorinini, Kallimini and Melitaeini, each of which is a well-supported, stable clade. In addition to the six tribes, there are several so-called "rogue" taxa (Kallimoides, Vanessula, Rhinopalpa and Pycina, all of which are monotypic), whose relationships are not entirely clear at the moment.


We have studied the evolutionary history of the subfamily in detail, mainly based on DNA sequences from three gene regions. The phylogenetic relationships of the 57 genera suggest that there are 10 major lineages in the subfamily (see Wahlberg et al. 2005, Wahlberg 2006). The subfamily appears to have diverged from it's sister group (possibly Cyrestinae) some 80 million years ago. The subfamily itself began diversifying after the catastrophic event that caused the mass extinction of e.g. the dinosaurs about 65 million years ago (the so-called K/T event). The first split was between the ancestors of the tribe Coeini and the rest of Nymphalinae, which appears to have happened soon after the K/T event. The timing of divergences in the rest of Nymphalinae has been studied in detail by Wahlberg (2006), and the results show that the deeper divergences in this group happened at the end of the Paleocene/beginning of the Eocene.


The tribe Coeini appears to contain only two genera: Historis and Baeotus. Historically the genera Colobura, Tigridia, Smyrna and Pycina have been associated with this tribe, but the former three are clearly within Nymphalini. Pycina has not been included in any published study, but recent unpublished molecular data generated by us is showing that the genus is a so-called "long-branch taxon" and it's position is not stable or well-supported in any analysis. Interestingly, Pycina appears to be sister to Rhinopalpa, another long-branch taxon. The four species of Baeotus form a well-supported, stable monophyletic group, whereas the two species of Historis do not. There may yet be cause to resurrect the genus Coea for the species acheronta if this turns out to be the sister group to Baeotus with more data. For the present, we prefer to retain it in the genus Historis. The tribe Coeini is entirely restricted to the New World, and the six species are mainly found in the tropical rainforests of South America. The position of the tribe as the sister group to the rest of Nymphalinae is being confirmed by a large dataset of sequences from 10 genes. Timings of divergences have not been studied for this group yet.


Vanessa cardui

Species in the tribe Nymphalini have been much studied in recent years. The tribe contains many well-known species, such as the red admiral and the painted lady. Temperate species are among the first species seen in Spring since they overwinter as adults. The phylogenetic relationships of the genera in the tribe are beginning to clear up. Colobura and Tigridia are sister genera and together appear to be sister to the rest of Nymphalini, although in some analyses Smyrna is sister to the two (see Wahlberg 2006). Whether Smyrna is sister to Colobura+Tigridia or to the rest of Nymphalini, both lineages appear to have diverged from the stem Nymphalini in the Paleocene some 60 to 55 million years ago. The genera Nymphalis, Aglais and Polygonia form a strongly supported and stable clade, known as the Nymphalis-group (Wahlberg and Nylin 2003), with Aglais being sister to the other two. The nomenclature of this group is contentious at the moment, with some people lumping the about 26 species into one genus Nymphalis, and others splitting the group into 8 or more genera. We prefer to retain three genera, although the species Polygonia canace is a difficult case that is sister to the rest of Polygonia only when morphological data is included in the analysis, perhaps justifying it's placement in a separate genus Kaniska. The genera Symbrenthia, Mynes and Araschnia also form a well-supported, stable monophyletic group, although the relationships of the three genera and their respective monophyly are not very clear at the moment (compare Fric et al. 2004 with Wahlberg et al. 2005). The relationships of the Nymphalis-group, Symbrenthia-group, Hypanartia, Antanartia and Vanessa are also still unstable and will require more data to resolve. Whatever the relationships are, it appears that the genera diverged from each other in the Eocene, between 50-35 million years ago.


Anartia lytrea

The recently resurrected tribe Victorinini was necessary when it was discovered that the tribe Kallimini of previous authors was paraphyletic with regard to Melitaeini (Wahlberg et al. 2005, Wahlberg 2006). Victorinini comprises the South American genera Metamorpha, Napeocles, Siproeta and Anartia. Relationships of the genera are a bit ambiguous, it appears that Napeocles is within Siproeta and that Metamorpha is either sister to Anartia, or sister to the rest of Victorinini. More data and taxa are clearly needed to resolve this. Relationships within Anartia appear to be clear (Blum et al. 2003). The genera in Victorinini appear to have diverged from each other in the Eocene about 50-40 million years ago.


Junoniini is another tribe resurrected based on recent results (Wahlberg et al. 2005, Wahlberg 2006). Junoniini is a largely African tribe with some species of Hypolimnas and Junonia making it to Asia and Australia, and a few species of Junonia to the New World. The genus Yoma is entirely restricted to Asia and Australia. Relationships of genera are quite robust with current data. Molecular results brought some surprises for this tribe. First of all, it is clear that Junonia and Precis (two genera that have been used interchangably for a couple of hundred years) are not each others sister groups, but rather Precis is sister to Hypolimnas. Junonia appears to be related to a clade containing the genera Protogoniomorpha and Yoma, with Salamis being sister to these three. The second surprise was that Salamis of many authors was not monophyletic, with species now in Protogoniomorpha being sister to Yoma. Thirdly, the two species in Kamilla are well within Junonia (thus Kamilla has been synonymised with Junonia), and it appears that the the species "Protogoniomorpha" cytora and "P." temora, belong most likely also in Junonia (Wahlberg et al. 2005). The tribe appears to have begun diversifying at the end of the Eocene/beginning of the Oligocene.


The reduced Kallimini comprises only 4 genera and 18 species. The relationships of the four genera are robust with current data, Kallima is sister to Catacroptera+Mallika, and Doleschallia is sister to these three. Doleschallia is largely restricted to the Australasian region, with one species coming in to SE Asia. Kallima, well-known for the near perfect leaf mimicking undersides of its wings, is restricted to tropical Asia, while Catacroptera and Mallika (both with single species) are found only in Africa.


Melitaea didyma

The tribe Melitaeini has been much studied ecologically and recently also phylogenetically (Kons 2000, Wahlberg & Zimmermann 2000, Zimmermann et al. 2000, Wahlberg et al. 2003, 2005, Wahlberg 2006, Wahlberg & Freitas 2007). The tribe has five major clades that could be named subtribes, the Euphydryas-group (Euphydryina), the Chlosyne-group, the Melitaea-group (Melitaeina), the Phyciodes-group (Phyciodina) and the Gnathotriche-group. Euphydryas is the sister group to the rest of Melitaeini with strong support, but the relationships of the other groups is not clear. Currently it looks like the Gnathotriche-group is sister to Phyciodina, with Melitaeina being sister to these two (Wahlberg et al. 2005, Wahlberg 2006, Wahlberg & Freitas 2007).

Incertae cedis genera

The relationships of the four species belonging to the genera Kallimoides, Vanessula, Rhinopalpa and Pycina are not clear at the moment. All four have very long branches based on molecular data, causing them to jump around in various analyses. All four are clearly part of Nymphalinae, and the first three appear to belong to the clade comprising Victorinini, Junoniini, Kallimini and Melitaeini, while Pycina is not settling down in any part of the Nymphalinae tree so far. Clearly more data are needed for these ancient taxa.

Key publications for Nymphalinae systematics:

  1. Achtelik, G. 1999: Vier neue arten aus der "Melitaea sultanensis-gruppe" (Staudinger, 1886) (Lepidoptera, Nymphalidae). - Atalanta 30: 63-86.
  2. Blum, M. J., E. Bermingham, and K. Dasmahapatra. 2003: A molecular phylogeny of the neotropical butterfly genus Anartia (Lepidoptera: Nymphalidae). Molecular Phylogenetics and Evolution 26:46-55.
  3. Field, W. D. 1971: Butterflies of the genus Vanessa and of the resurrected genera Bassaris and Cynthia (Lepidoptera: Nymphalidae). - Smithsonian Contributions to Zoology 84: 1-105.
  4. Freitas, A. V. L. 1991: Variation, life cycle and systematics of Tegosa claudina (Eschscholtz) (Lepidoptera, Nymphalidae, Melitaeinae) in São Paulo state, Brazil. - Revta bras. Ent. 35: 301-306.
  5. Fric, Z., Konvicka, M. & Zrzavy, J. 2004: Red & black or black & white? Phylogeny of the Araschnia butterflies (Lepidoptera: Nymphalidae) and evolution of seasonal polyphenism. - Journal of Evolutionary Biology 17:265-278.
  6. Fox, R. M. & Forbes, A. C. 1971: The butterflies of the genera Siproeta and Metamorpha (Lepidoptera: Nymphalidae). - Annals of Carnegie Museum 43: 223-247.
  7. Higgins, L. G. 1941: An illustrated catalogue of the Palearctic Melitaea (Lep. Rhopalocera). - Trans. R. Ent. Soc. Lond. 91: 175-365.
  8. Higgins, L. G. 1950: A descriptive catalogue of the Palearctic Euphydryas (Lepidoptera: Rhopalocera). - Trans. R. Ent. Soc. Lond. 101: 435-487.
  9. Higgins, L. G. 1955: A descriptive catalogue of the genus Mellicta Billberg (Lepidoptera: Nymphalidae) and its species, with supplementary notes on the genera Melitaea and Euphydryas. - Trans. R. Ent. Soc. Lond. 10: 61-127.
  10. Higgins, L. G. 1960: A revision of the melitaeine genus Chlosyne and allied species (Lepidoptera: Nymphalinae). - Trans. R. Ent. Soc. Lond. 112: 381-465.
  11. Higgins, L. G. 1978: A revision of the genus Euphydryas Scudder (Lepidoptera: Nymphalidae). - Entomologist's Gaz. 29: 109-115.
  12. Higgins, L. G. 1981: A revision of Phyciodes Hübner and related genera, with a review of the classification of the Melitaeinae (Lepidoptera: Nymphalidae). - Bull. Brit. Mus. Nat. Hist. 43: 77-243.
  13. Kolesnichenko, K. A. 1999: A review of the East-Palaearctic taxa of the Melitaea didyma (Esper, 1779)-group 1. The M. ala Staudinger, 1881-M. chitralensis Moore, 1901-complex (Lepidoptera, Nymphalidae). - Atalanta 30: 87-117.
  14. Kons, H. L. Jr. 2000: Phylogenetic studies of the Melitaeini (Lepidoptera: Nymphalidae: Nymphalinae) and a revision of the genus Chlosyne Butler. - PhD thesis, University of Florida, Gainesville, 799 pp. Available from UMI.
  15. Larsen, T. B. 1991: The Butterflies of Kenya and their Natural History. - Oxford University Press, Oxford, UK.
  16. Nylin, S., K. Nyblom, F. Ronquist, N. Janz, J. Belicek, and M. Källersjö. 2001. Phylogeny of Polygonia, Nymphalis and related butterflies (Lepidoptera: Nymphalidae): a total-evidence analysis. Zoological Journal of the Linnean Society 132:441-468.
  17. Scott, J. A. 1994: Biology and systematics of Phyciodes (Phyciodes). - Papilio (New Series) 7: 1-120 (privately published, available from author).
  18. Scott, J. A. 1998: Phyciodes (Phyciodes): new discoveries, new subspecies, and convergence. - Papilio (New Series) 10: 1-42 (privately published, available from author).
  19. Shirôzu, T. and Nakanishi, A. 1984: A revision of the genus Kallima Doubleday (Lepidoptera, Nymphalidae). - Tyo to Ga 34: 97-110.
  20. Silberglied, R. E., Aiello, A. & Lamas, G. 1979: Neotropical butterflies of the genus Anartia: systematics, life histories and general biology (Lepidoptera: Nymphalidae). - Psyche 86: 219-260.
  21. Wahlberg, N. 2006. That awkward age for butterflies: insights from the age of the butterfly subfamily Nymphalinae. Systematic Biology 55:703-714.
  22. Wahlberg, N., A. V. Z. Brower, and S. Nylin. 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.
  23. Wahlberg, N., and A. V. L. Freitas. 2007. Colonization of and radiation in South America by butterflies in the subtribe Phyciodina (Lepidoptera: Nymphalidae). Molecular Phylogenetics and Evolution in press.
  24. Wahlberg, N., and S. Nylin. 2003. Morphology versus molecules: resolution of the positions of Nymphalis, Polygonia and related genera (Lepidoptera: Nymphalidae). Cladistics 19:213-223.
  25. Wahlberg, N., R. Oliveira, and J. A. Scott. 2003. Phylogenetic relationships of Phyciodes butterfly species (Lepidoptera: Nymphalidae): complex mtDNA variation and species delimitations. Systematic Entomology 28:257-273.
  26. Wahlberg, N., E. Weingartner, and S. Nylin. 2003. Towards a better understanding of the higher systematics of Nymphalidae (Lepidoptera: Papilionoidea). Molecular Phylogenetics and Evolution 28:473-484.
  27. Wahlberg, N., and M. Zimmermann. 2000. Pattern of phylogenetic relationships among members of the tribe Melitaeini (Lepidoptera: Nymphalidae) inferred from mtDNA sequences. Cladistics 16:347-363.
  28. Weingartner, E., N. Wahlberg, and 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.
  29. Weiss, D. & Major, V. 2000: A new species of the genus Melitaea Fabricius, 1807 from Iran. - Atalanta 31: 123-128.
  30. Willmott, K. R., Constantino, L. M. & Hall, J. P. W. 2001a: A review of Colobura (Lepidoptera: Nymphalidae) with comments on larval and adult ecology and description of a sibling species. - Ann. Entomol. Soc. Am. 94: 185-196.
  31. Willmott, K. R., Hall, J. P. W. & Lamas, G. 2001b: Systematics of Hypanartia (Lepidoptera: Nymphalidae: Nymphalinae), with a test for geographical speciation mechanisms in the Andes. - Systematic Entomology 26: 369-399.
  32. Zimmermann, M., N. Wahlberg, and H. Descimon. 2000. Phylogeny of Euphydryas checkerspot butterflies (Lepidoptera: Nymphalidae) based on mitochondrial DNA sequence data. Annals of the Entomological Society of America 93:347-355.