Noctuoidea

Noctuoidea
File:Noctua.pronuba.7199.jpg
Large Yellow Underwing, Noctua pronuba
Scientific classification
Kingdom:	Animalia
Phylum:	Arthropoda
Class:	Insecta
Order:	Lepidoptera
Suborder:	Glossata
Infraorder:	Heteroneura
(unranked):	Ditrysia
(unranked):	Macrolepidoptera
Superfamily:	Noctuoidea
Families
Erebidae Noctuidae strict sense Nolidae Notodontidae Oenosandridae Doidae (?) (See text)
Diversity
over 70,000 species

Noctuoidea is the superfamily of noctuid (Latin "night owl") or "owlet" moths, and has more than 70000 described species, the largest number of for any Lepidopteran superfamily. Its classification has not yet reached a satisfactory or stable state. Since the end of the 20th century, increasing availability of molecular phylogenetic data for this hugely successful radiation has led to several competing proposals for a taxonomic arrangement that correctly represents the relationships between the major lineages.

Briefly, the disputes center on the fact that in old treatments (which were just as unable to reach a general consensus) the distinctness of some groups, such as the Arctiidae or Lymantriidae, was overrated due to their characteristic appearance, while some less-studied lineages conventionally held to be Noctuidae are in fact quite distinct. This requires a rearrangement at least of the latter family (by simply including anything disputed within it). This is quite unwieldy, and various more refined treatments have been proposed in response to it. While there is general agreement on what the basal families of Noctuoidea are, the more diverse advanced group may be treated as one all-encompassing Noctuidae, two huge and two smaller, or even (if Arctiidae or Lymantriida are kept distinct) more than four families, which are in some cases still quite sizeable.

Recent developments

There are several recent studies suggesting a radical change in the traditional family level classification. Recent works in redifining the families within the Noctuoidea has been carried out by Kitching (1984), Poole (1995), Kitching and Rawlins [1998], Speidel et al. (1996), Mitchell et al. (1997, 2000, 2006), Fibiger and Lafontaine (2005), Lafontaine and Fibiger (2006) and Zahiri et al. (2010).

The Noctuoidea can be divided into two broad groups, those with a trifid forewing venation (Oenosandridae, Notodontidae and Doidae), and those with a quadrifid forewing venation (e.g., Arctiidae, Lymantriidae, Nolidae, Noctuidae). What has emerged from these investigations is that the quadrifid Noctuoidea form a monophyletic group. In 2005, Fibiger and Lafontaine arranged the quadrifid (forewing) group into several families, including the quadrifine (hindwing) Erebidae and trifine (hindwing) Noctuidae, based on evidence that suggested that the trifine noctuid subfamilies were derived from within the quadrifine subfamilies, so the family Erebidae would not be strictly monophyletic.

Lafontaine and Fibiger in 2006 then redefined the Noctuidae to include the entire quadrifid group, believing the Arctiidae, Lymantriidae, and Nolidae to be derived from within this expanded concept of Noctuidae (and closely related to the subfamily Catocalinae). In essence, groups such as the Arctiidae, which had previously been treated as a separate family, were more closely related to groups within the Noctuidae than to non-noctuid families. In order to address this, a revised classification would have meant either recognizing over 20 (often weakly defined) families, or a single well-defined family with numerous subfamilies. The latter was adopted (Lafontaine and Fibiger 2006).

More recent evidence from nuclear genes (Zahiri et al. 2010) confirms that the quadrifid (forewing) noctuoids form a monophyletic group, but also that this group can be further arranged into four monophyletic subgroups: 1) the quadrifine subfamilies; 2) the trifine subfamilies; 3) the Nolinae; and 4) the Euteliinae. Considering the massive size of the family, and the large number of subfamilies, tribes, and subtribes to arrange into a classification, Zahiri et al. (2010) chose the option of recognizing these four groups as families, namely Erebidae, Noctuidae, Nolidae, and Euteliidae, in addition to the basal trifid families.

Systematics

This follows Lafontaine & Fibiger (2006), with the additions of Thiacidinae Hacker & Zilli, 2007 and taxa in Micronoctuidae Fibiger, 2005. Note that the placement of Arctiidae, Lymantriidae and Nolidae as subfamilies of Noctuidae has been largely rejected by subsequent authors. An updated classification for North American Noctuoidea has recently been published (Lafontaine & Schmidt, 2010) and further changes are imminent for a global molecular review of the superfamily (Zahiri et al., in press).

• Familia Oenosandridae Miller, 1991
• Familia Doidae Donahue & Brown, 1987
• Familia Notodontidae Stephens, 1829
  • Subfamilia Thaumetopoeinae Aurivillius, 1889
  • Subfamilia Pygaerinae Duponchel, 1845
  • Subfamilia Platychasmatinae Nakamura, 1956
  • Subfamilia Notodontinae Stephens, 1829
    • Tribus Notodontini Stephens, 1829
    • Tribus Dicranurini Duponchel, 1845
  • Subfamilia Phalerinae Butler, 1886
  • Subfamilia Dudusinae Matsumura, 1925
    • Tribus Dudusini Matsumura, 1925
    • Tribus Scranciini Miller, 1991
  • Subfamilia Hemiceratinae Guenée, 1852
  • Subfamilia Heterocampinae Neumogen & Dyar, 1894
  • Subfamilia Nystaleinae Forbes, 1948
  • Subfamilia Dioptinae Walker, 1862
• Familia Micronoctuidae Fibiger, 2005
  • Subfamilia Micronoctuinae Fibiger, 2005
  • Subfamilia Pollexinae Fibiger, 2007
  • Subfamilia Belluliinae Fibiger, 2008
    • Tribus Belluliini Fibiger, 2008
    • Tribus Medialini Fibiger, 2008
  • Subfamilia Magninae Fibiger, 2008
    • Tribus Magnini Fibiger, 2008
    • Tribus Faeculini Fibiger, 2008
  • Subfamilia Parachrostiinae Fibiger, 2008
    • Tribus Duplexini Fibiger, 2008
    • Tribus Parachrostiini Fibiger, 2008
• Familia Noctuidae Latreille, 1809
  • Subfamilia Rivulinae Grote, 1895
  • Subfamilia Boletobiinae Grote, 1895
  • Subfamilia Hypenodinae Forbes, 1954
  • Subfamilia Araeopteroninae Fibiger, 2005
  • Subfamilia Eublemminae Forbes, 1954
    • Tribus Eublemmini Forbes, 1954
    • Tribus Pangraptini Grote, 1882
  • Subfamilia Herminiinae Leach, 1815
  • Subfamilia Scolecocampinae Grote, 1883
  • Subfamilia Hypeninae Herrich-Schäffer, 1851
  • Subfamilia Phytometrinae Hampson, 1913
  • Subfamilia Aventiinae Tutt, 1896
  • Subfamilia Erebinae Leach, 1815
  • Subfamilia Calpinae Boisduval, 1840
    • Tribus Anomini Grote, 1882
    • Tribus Calpini Boisduval, 1840
    • Tribus Phyllodini Guenée, 1852
  • Subfamilia Catocalinae Boisduval, 1828
    • Tribus Toxocampini Guenée, 1852
    • Tribus Acantholipini Fibiger & Lafontaine, 2005
    • Tribus Arytrurini Fibiger & Lafontaine, 2005
    • Tribus Melipotini Grote, 1895
    • Tribus Euclidiini Guenée, 1852
    • Tribus Panopodini Forbes, 1954
    • Tribus Ophiusini Guenée, 1837 (= Omopterini Boisduval, 1833, suppressed older syn.)
    • Tribus Catocalini Boisduval, 1828
    • Tribus Anobini Holloway, 2005 (= Anobini Wiltshire, 1990, nomen nudum)
    • Tribus Sypnini Holloway, 2005
    • Tribus Hypopyrini Guenée, 1852
    • Tribus Tinolini Moore, 1885
    • Tribus Hulodini Guenée, 1852 (= Speiredoniinae Swinhoe, 1900)
    • Tribus Ommatophorini Guenée, 1852
    • Tribus Pericymini Wiltshire; 1976
    • Tribus Pandesmini Wiltshire, 1990, nomen nudum
    • Tribus Catephiini Guenée, 1852
    • Tribus Ercheini Berio, 1992
  • Subfamilia Cocytiinae Boisduval, 1874
  • Subfamilia Stictopterinae Hampson, 1894
  • Subfamilia Euteliinae Grote, 1882
  • (Subfamilia Nolinae) Bruand, 1846, stat rev.
    • Tribus Nolini Bruand, 1846, stat rev.
    • Tribus Chloephorini Stainton, 1859, stat rev.
      • Subtribus Chloephorina Stainton, 1859, stat rev.
      • Subtribus Sarrothripina Hampson, 1894, stat rev.
      • Subtribus Camptolomina Mell, 1943, stat rev.
      • Subtribus Careina Moore, 1883, stat rev.
      • Subtribus Ariolicina Mell, 1943, stat rev.
    • Tribus Westermanniini Hampson, 1918, stat rev.
    • Tribus Eariadini Hampson, 1912, stat rev.
    • Tribus Blenini Mell, 1943, stat rev.
    • Tribus Risobini Mell, 1943, stat rev.
    • Tribus Collomenini Kitching & Rawlins, 1998, stat rev.
    • Tribus Afridini Kitching & Rawlins, 1998, stat rev.
    • Tribus Eligmini Mell, 1943, stat rev.
  • Subfamilia Aganainae Boisduval, 1833
  • (Subfamilia Arctiinae Leach, 1815), stat. nov.
    • Tribus Lithosiini Billberg, 1820, stat. rev.
      • Subtribus Phryganopterygina Bendib & Minet, 1999, stat. rev.
      • Subtribus Aesalina Bendib & Minet, 1999, stat. rev.
      • Subtribus Eudesmiina Bendib & Minet, 1999, stat. rev.
      • Subtribus Cisthenina Bendib & Minet, 1999, stat. rev.
      • Subtribus Nudariina Börner, 1920, stat. rev.
      • Subtribus Endrosina Börner, 1932, stat. rev.
      • Subtribus Lithosiina Billberg, 1820, stat. rev.
    • Tribus Syntomini Herrich-Schäffer, 1846, stat. rev.
      • Subtribus Syntomina Herrich-Schäffer, 1846, stat. rev.
      • Subtribus Thyretina Butler, 1876, stat. rev.
    • Tribus Arctiini Leach, 1815, stat. rev.
      • Subtribus Arctiina Leach, 1815, stat. rev.
      • Subtribus Callimorphina Walker, 1865, stat. rev.
      • Subtribus Pericopina Walker, 1865, stat. rev.
      • Subtribus Phaegopterina Kirby, 1892, stat. rev.
      • Subtribus Ctenuchina Kirby, 1837, stat. rev.
      • Subtribus Euchromiina Butler, 1876, stat. rev.
  • (Subfamilia Lymantriinae) Hampson, 1893, stat. nov.
    • Tribus Lymantriini Hampson, 1893, stat. rev.
    • Tribus Orgyiini Wallengren, 1861, stat. rev.
    • Tribus Arctornithini Holloway, 1999, stat. rev.
    • Tribus Leucomini Grote, 1895, stat. rev.
    • Tribus Nygmiini Holloway, 1999, stat. rev.
  • Subfamilia Strepsimaninae Meyrick, 1930, stat rev.
  • Subfamilia Plusiinae Boisduval, 1828
    • Tribus Abrostolini Eichlin & Cunningham, 1978
    • Tribus Argyrogrammatini Eichlin & Cunningham, 1978
    • Tribus Plusiini Boisduval, 1828
      • Subtribus Autoplusiina Kitching, 1987
      • Subtribus Euchaciina Chou & Lu, 1979
      • Subtribus Plusiina Boisduval, 1828
  • Subfamilia Eustrotiinae Grote, 1882
  • Subfamilia Bagisarinae Crumb, 1956
    • Tribus Bagisarini Crumb, 1956
    • Tribus Cydosiini Kitching & Rawlins, 1998
  • Subfamilia Acontiinae Guenée, 1841
    • Tribus Hypercalymniini Fibiger & Lafontaine, 2005
    • Tribus Acontiini Guenée, 1841
    • Tribus Armadini Wiltshire, 1961
    • Tribus Aediini Beck, 1960
  • Subfamilia Pantheinae Smith, 1898
  • Subfamilia Diphtherinae Fibiger & Lafontaine, 2005
  • Subfamilia Dilobinae Aurivillius, 1889
  • Subfamilia Raphiinae Beck, 1996
  • Subfamilia Balsinae Grote, 1896, stat. rev.
  • Subfamilia Acronictinae Heinemann, 1859
  • Subfamilia Metoponiinae Herrich-Schäffer, 1851
  • Subfamilia Sinocharinae Speidel, Fänger & Naumann, 1996
  • Subfamilia Lophonyctinae Speidel, Fänger & Naumann, 1996
  • Subfamilia Agaristinae Herrich-Schäffer, 1858
  • Subfamilia Eucocytiinae Hampson, 1918
  • Subfamilia Cuculliinae Herrich-Schäffer, 1850
  • Subfamilia Oncocnemidinae Forbes & Franclemont, 1954
  • Subfamilia Amphipyrinae Guenée, 1837
  • Subfamilia Psaphidinae Grote, 1896
    • Tribus Psaphidini Grote, 1896
    • Tribus Feraliini Poole, 1995
    • Tribus Nocloini Poole, 1995
    • Tribus Triocnemidini Poole, 1995
  • Subfamilia Stiriinae Grote, 1882
    • Tribus Stiriini Grote, 1882
    • Tribus Grotellini Poole, 1995
    • Tribus Azenini Poole, 1995
  • Subfamilia Heliothinae Boisduval, 1828
  • Subfamilia Condicinae Poole, 1995
    • Tribus Condicini Poole, 1995
    • Tribus Leuconyctini Poole, 1995
  • Subfamilia Eriopinae Herrich-Schäffer, 1851
  • Subfamilia Bryophilinae Guenée, 1852
  • Subfamilia Xyleninae Guenée, 1837
    • Tribus Pseudeustrotiini Beck, 1996
    • Tribus Phosphilini Poole, 1995, stat. rev.
    • Tribus Prodenilni Forbes, 1954
    • Tribus Eiaphriini Beck, 1996
    • Tribus Caradrinini Boisduval, 1840
      • Subtribus Caradrinina Boisduval, 1840
      • Subtribus Athetina Fibiger & Lafontaine, 2005
    • Tribus Dypterygiini Forbes, 1954
    • Tribus Actinotiini Beck, 1996
    • Tribus Phlogophorini Hampson, 1918
    • Tribus Apameini Guenée, 1841 (= Nonagriini Guenée, 1837, suppressed older syn.)
      • Subtribus Oxytripiina Gozmany, 1970
      • Subtribus Apameina Guenée, 1841
      • Subtribus Sesamiina Fibiger & Goldstein, 2005
    • Tribus Arzamini Grote, 1883
    • Tribus Episemini Guenée, 1852
    • Tribus Xylenini Guenée, 1837
      • Subtribus Xylenina Guenée, 1837
      • Subtribus Cosmiina Guenée, 1852, stat. rev.
      • Subtribus Antitypina Forbes & Franclemont, 1954
      • Subtribus Ufeina Crumb, 1956
  • Subfamilia Hadeninae Guenée, 1837
    • Tribus Orthosiini Guenée, 1837
    • Tribus Tholerini Beck, 1996
    • Tribus Hadenini Guenée, 1837
    • Tribus Leucaniini Guenée, 1837
    • Tribus Eriopygini Fibiger & Lafontaine, 2005
    • Tribus Glottulini Guenée, 1852
  • Subfamilia Noctuinae Latreille, 1809
    • Tribus Agrotini Rambur, 1848
      • Subtribus Austrandesiina Angulo & Olivares, 1990
      • Subtribus Agrotina Rambur, 1848
    • Tribus Noctuini Latreille, 1809
      • Subtribus Axyliina Fibiger & Lafontaine, 2005
      • Subtribus Noctuina Latreille, 1809

References

• Fibiger, M., 2007. Revision of the Micronoctuidae (Lepidoptera: Noctuoidea). Part 1, Taxonomy of the Pollexinae. Zootaxa 1567: 1–116. (abstract)
• Fibiger, M., 2008. Revision of the Micronoctuidae (Lepidoptera: Noctuoidea). Part 2, Taxonomy of the Belluliinae, Magninae, and Parachrostiinae. Zootaxa 1867: 1–136. (abstract)
• Hacker, H.H. & Zilli, A., 2007. Esperiana Buchreihe zur Entomologie Memoir 3: 179-246.
• Kitching, I.J. & Rawlins, J.E., 1999. The Noctuoidea. In: Lepidoptera, Moths and Butterflies, Volume 1: Evolution, Systematics, and Biogeography, ed. N. P.Kristensen, pp. 355-401. Walter de Gruyter, Berlin.
• Lafontaine, J.D. & Fibiger, M., 2006. Revised higher classification of the Noctuoidea (Lepidoptera). Canadian Entomologist 138(5):610-635 (abstract).
• Lafontaine, J.D. & Schmidt, C., 2010. Annotated check list of the Noctuoidea (Insecta, Lepidoptera) of North America north of Mexico. ZooKeys 40: 1-239.
• O'Toole, C. (ed.), 2002. Firefly Encyclopedia of Insects and Spiders. ISBN 1-55297-612-2.
• Zahiri, R., Kitching, I.J., Lafontaine, J.D., Mutanen, M., Kaila, L., Holloway, J.D. & Wahlberg, N. (in press) A new molecular phylogeny offers hope for a stable family-level classification of the Noctuoidea (Insecta: Lepidoptera). Molecular Phylogenetics and Evolution.

External links

• Tree of Life
• Noctuoidea (summary of R.Zahiri's ongoing Ph.D. studies)