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Order Lepidoptera Linnaeus, 1758. In: Zhang Z-Q, editor. 2011. Animal
Biodiversity: An outline of higher classification and survey of taxonomic
richness

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Order Lepidoptera Linnaeus, 1758 (4 suborders) (15,578 genera, 157,424 species, 50/86) 1 2 3
ಲUnassigned early lepidopterans (4 families)
ಲFamily unassigned (12 genera, 16 species, 12/16) ϰ
ಲ Family Archaeolepidae Whalley, 1985 (1 genus, 1 species, 1/1) ϱ
ಲ Family Mesokristenseniidae Huang, Nel & Minet, 2010 (1 genus, 3 species, 1/3) ϲ
ಲ Family Eolepidopterigidae Rasnitsyn, 1983 (1 genus, 1 species, 1/1) ϳ
ಲ Family Undopterigidae Kozlov, 1988 (1 genus, 1 species, 1/1) ϴ
Suborder Zeugloptera Chapman, 1917 (1 superfamily)
Superfamily Micropterigoidea Herrich-Schäffer, 1855 (1 family)
Family Micropterigidae Herrich-Schäffer, 1855 (21 genera, 160 species, 3/6) 9
Suborder Aglossata Speidel, 1977 (1 superfamily)
Superfamily Agathiphagoidea Kristensen, 1967 (1 family)
Family Agathiphagidae Kristensen, 1967 (1 genus, 2 species)
Suborder Heterobathmiina Kristensen & Nielsen, 1983 (1 superfamily)
Superfamily Heterobathmioidea Kristensen & Nielsen, 1979 (1 family)
Family Heterobathmiidae Kristensen & Nielsen, 1979 (1 genus, 3 species)
Suborder Glossata Fabricius, 1775 (6 infraorders, all following)
Infraorder Dacnonypha Hinton, 1946 (1 superfamily)
Superfamily Eriocranioidea Rebel, 1901 (1 family)
Family Eriocraniidae Rebel, 1901 (5 genera, 29 species, 0/1)
Clade Coelolepida Nielsen & Kristensen, 1996 (5 infraorders, all following) 10
Infraorder Acanthoctesia Minet, 2002 (1 superfamily)
Superfamily Acanthopteroctetoidea Davis, 1978 (1 family)
Family Acanthopteroctetidae Davis, 1978 (2 genera, 5 species)
Infraorder Lophocoronina Common, 1990 (1 superfamily)
Superfamily Lophocoronoidea Common, 1973 (1 family)
Family Lophocoronidae Common, 1973 (1 genus, 6 species)
Clade Myoglossata Kristensen & Nielsen, 1981 (3 infraorders, all following)
Infraorder Neopseustina Davis & Nielsen, 1980 (1 superfamily)
Superfamily Neopseustoidea Hering, 1925 (1 family)
Family Neopseustidae Hering, 1925 (4 genera, 14 species)
Clade Neolepidoptera Packard, 1895 (2 infraorders, all following)

1. By Erik J. van Nieukerken, Lauri Kaila, Ian J. Kitching, Niels P. Kristensen, David C. Lees, Joël Minet, Charles Mitter, Marko Mutanen, Jerome
C. Regier, Thomas J. Simonsen, Niklas Wahlberg, Shen-Horn Yen, Reza Zahiri, David Adamski, Joaquin Baixeras, Daniel Bartsch, Bengt Å.
Bengtsson, John W. Brown, Sibyl Rae Bucheli, Donald R. Davis, Jurate De Prins, Willy De Prins, Marc E. Epstein, Patricia Gentili-Poole, Cees
Gielis, Peter Hättenschwiler, Axel Hausmann, Jeremy D. Holloway, Axel Kallies, Ole Karsholt, Akito Y. Kawahara, Sjaak (J.C.) Koster,
Mikhail V. Kozlov, J. Donald Lafontaine, Gerardo Lamas, Jean-François Landry, Sangmi Lee, Matthias Nuss, Kyu-Tek Park, Carla Penz,
Jadranka Rota, Alexander Schintlmeister, B. Christian Schmidt, Jae-Cheon Sohn, M. Alma Solis, Gerhard M. Tarmann, Andrew D. Warren,
Susan Weller, Roman V. Yakovlev, Vadim V. Zolotuhin, Andreas Zwick (for full contact details, see the list after References). The title of this
contribution should be cited as “Order Lepidoptera Linnaeus, 1758. In: Zhang, Z.-Q. (Ed.) Animal biodiversity: An outline of higher-level
classification and survey of taxonomic richness”.
2. The classification largely follows that in the Handbook of Zoology (Kristensen 1998), and the later update (Kristensen et al. 2007). Recent
molecular studies of Ditrysia (Regier et al. 2009; Mutanen et al. 2010) are responsible for several novelties, such as the position of the butterflies
(Papilionoidea) and the synonymy of Sesioidea with Cossoidea.
3. When possible, numbers of genera and species are based on counts in existing databases, whether published (then cited) or personal. Where not
available, often for larger taxa, numbers provided by Handbook authors were updated with Zoological Record data (up to early July 2011).
Lepindex (Beccaloni et al. 2005) has been an additional source; x/y for numbers of genera/species represented by fossil.
4. Including "homoneurous" (or probably homoneurous) lepidopteran genera not certainly placed to family (taxa listed in Table 13.1 [1, 3-6] of
Grimaldi & Engel (2005), plus - at least - the following six genera: Archiptilia Handlirsch, 1939, Electrocrania Kusnezov, 1941, Epididontus
Handlirsch, 1939, Palaeosabatinca Kozlov, 1988, Parataulius Handlirsch, 1939, and Xena Martins-Neto, 1999).
5. The presence of wing scales is supposed to indicate that this fossil is lepidopteran (Kristensen & Skalski 1998; Grimaldi & Engel 2005). This
family and the following three are listed in "stratigraphical" order.
6. A group from the middle Jurassic of China best placed among unassigned early Lepidoptera, although this group might be sister to
Micropterigidae (Huang et al. 2010).
7. Including with certainty only Eolepidopterix jurassica Rasnitsyn, 1983.
8. Including with certainty only Undopterix sukatshevae Skalski, 1979.
9. With the already known, but still unnamed species, the global total is around 260 species.
10. Junior homonym of Coelolepida in Agnatha, but unlikely to be confused.

212 · Zootaxa 3148 © 2011 Magnolia Press VAN NIEUKERKEN ET AL.: ORDER LEPIDOPTERA
 Infraorder Exoporia Common, 1975 (2 superfamilies) 11
Superfamily Mnesarchaeoidea Eyer, 1924 (1 family)
Family Mnesarchaeidae Eyer, 1924 (1 genus, 7 species)
Superfamily Hepialoidea Stephens, 1829 (5 families)
Family Palaeosetidae Turner, 1922 (4 genera, 9 species)
Family Prototheoridae Meyrick, 1917 (1 genus, 12 species)
Family Neotheoridae Kristensen, 1978 (1 genus, 1 species)
Family Anomosetidae Tillyard, 1919 (1 genus, 1 species)
Family Hepialidae Stephens, 1829 (62 genera, 606 species, 2/2)
Infraorder Heteroneura Tillyard, 1918 (34 superfamilies, all following)
Clade Nepticulina Meyrick, 1928 (1 superfamily)
Superfamily Nepticuloidea Stainton, 1854 (2 families) 12
Family Nepticulidae Stainton, 1854 (13 genera, 819 species, 1/13)
Family Opostegidae Meyrick, 1893 (7 genera, 192 species)
Clade Eulepidoptera Kiriakoff, 1948 (33 superfamilies, all following)
Clade Incurvariina Börner, 1939 (2 superfamilies)
Superfamily Andesianoidea Davis & Gentili, 2003 (1 family)
Family Andesianidae Davis & Gentili, 2003 (1 genus, 3 species)
Superfamily Adeloidea Bruand, 1850 (5 families) 13
Family Heliozelidae Heinemann & Wocke, 1876 (12 genera, 123 species)
Family Adelidae Bruand, 1850 (5 genera, 294 species)
Family Incurvariidae Spuler, 1898 (11 genera, 51 species) 14
Family Cecidosidae Bréthes, 1916 (5 genera, 16 species)
Family Prodoxidae Riley, 1881 (9 genera, 98 species)
Clade Etimonotrysia Minet, 1984 (2 superfamilies)
Superfamily Palaephatoidea Davis, 1986 (1 family)
Family Palaephatidae Davis, 1986 (7 genera, 57 species)
Superfamily Tischerioidea Spuler, 1898 (1 family)
Family Tischeriidae Spuler, 1898 (3 genera, 110 species) 15
Clade Ditrysia Börner, 1925 (29 superfamilies, all following)
Superfamily unassigned (1 family)
Family unassigned (25 genera, 100 species) 16
Family Millieriidae Heppner, 1982 (3 genera, 4 species) 17
Superfamily Tineoidea Latreille, 1810 (3 families) 18
Family Eriocottidae Spuler, 1898 (6 genera, 80 species)
Family Psychidae Boisduval, 1829 (241 genera, 1,350 species) 19
Family Tineidae Latreille, 1810 (357 genera, 2,393 species) 20
Superfamily Gracillarioidea Stainton, 1854 (3 families) 21

11. Diversity of all families in Exoporia updated from Nielsen et al. (2000).
12. See van Nieukerken (2010).
13. The frequently used name Incurvarioidea is a junior synonym of Adeloidea. As the latter name has been used occasionally before 2000, ICZN
article 35.5 does not apply, and priority has to be followed here.
14. The monobasic Crinopterygidae Spuler, 1898 is here considered a subfamily of Incurvariidae.
15. Updated from Diškus & Puplesis (2003).
16. This group comprises a number of genera that cannot be placed with confidence in any currently known family, although some could be
attributed to one of the following superfamilies. New family group taxa are probably needed for some of these. Included here are, e.g.: the
probably non-monophyletic Pseudurgis group, excluded from Cossidae (Mey 2007), the Heliocosma group, excluded from Tortricidae (Horak
& Common 1996), the genus Titanomis Meyrick, 1888 (Hoare 2001), the genus Allotoma Roepke, 1944 (Holloway 1999), which is probably
Bombycoidea, a number of genera excluded from Heliodinidae (Hsu & Powell 2005), Kenguichardia Holloway, 1987 in Noctuoidea, and
several others.
17. Raised to family by Rota (2011).
18. Although Tineoidea could not be shown to be monophyletic (Mutanen et al. 2010), we retain it as support is conflicting for the dismembering of
the superfamily.
19. Psychidae now also contain Arrhenophaninae as a subfamily (Mutanen et al. 2010). Diversity updated with Arrhenophanidae from Sobczyk
(2011).
20. See Robinson (2009). Acrolophinae are included as a subfamily of Tineidae (Mutanen et al. 2010).

ANIMAL BIODIVERSITY Zootaxa 3148 © 2011 Magnolia Press · 213

 Family Roeslerstammiidae Bruand, 1850 (13 genera, 53 species) 22
Family Bucculatricidae Fracker, 1915 (4 genera, 297 species) 23
Family Gracillariidae Stainton, 1854 (101 genera, 1,866 species, 1/2) 24
Superfamily Yponomeutoidea Stephens, 1829 (10 families)
Family Yponomeutidae Stephens, 1829 (95 genera, 363 species) 25
Family Argyresthiidae Bruand, 1850 (1 genus, 157 species)
Family Plutellidae Guenée, 1845 (48 genera, 150 species)
Family Glyphipterigidae Stainton, 1854 (28 genera, 535 species) 26
Family Ypsolophidae Guenée, 1845 (7 genera, 163 species)
Family Attevidae Mosher, 1916 (1 genus, 52 species)
Family Praydidae Moriuti, 1977 (3 genera, 47 species)
Family Heliodinidae Heinemann & Wocke, 1876 (13 genera, 69 species)
Family Bedelliidae Meyrick, 1880 (1 genus, 16 species)
Family Lyonetiidae Stainton, 1854 (32 genera, 204 species)
Clade Apoditrysia Minet, 1983 (26 superfamilies, all following) 27
Superfamily unassigned (2 families)
Family Prodidactidae Epstein & Brown, 2003 (1 genus, 1 species)
Family Douglasiidae Heinemann & Wocke, 1876 (2 genera, 29 species) 28
Superfamily Simaethistoidea Minet, 1991 (1 family)
Family Simaethistidae Minet, 1991 (2 genera, 4 species)
Superfamily Gelechioidea Stainton, 1854 (21 families) 29 30
Family Autostichidae Le Marchand, 1947 (72 genera, 638 species) 31
Family Lecithoceridae Le Marchand, 1947 (100 genera, 1,200 species)
Family Xyloryctidae Meyrick, 1890 (60 genera, 524 species)
Family Blastobasidae Meyrick, 1894 (24 genera, 377 species) 32
Family Oecophoridae Bruand, 1850 (313 genera, 3,308 species) 33
Family Schistonoeidae Hodges, 1998 (1 genus, 1 species)
Family Lypusidae Herrich-Schäffer, 1857 (3 genera, 21 species) 34
Family Chimabachidae Heinemann, 1870 (2 genera, 6 species)
Family Peleopodidae Hodges, 1974 (7 genera, 28 species) 35

21. Although Roeslerstammiidae is older than Gracillariidae, Gracillarioidea can be used as superfamily name, following ICZN article 35.5:
Roeslerstammioidea has never been used as a published name prior to 2000.
22. Amphitheridae Meyrick, 1914 is a junior synonym: see van Nieukerken & Karsholt (2006).
23. Tritymba Lower, 1894 (with nine Australian species), is transferred from Plutellidae to Bucculatricidae following the results of the analysis by
Mutanen et al. (2010).
24. See De Prins & De Prins (2011a).
25. Since the monophyly of the extended Yponomeutidae in the sense of Kyrki (1990) had already been strongly questioned (e.g. Dugdale et al.
1998), and in molecular analyses (Mutanen et al. 2010, C. Mitter pers. obs.) constituent taxa never form a monophylum, we have placed
Argyresthiidae, Praydidae and Attevidae as separate families.
26. This now includes subfamilies Orthoteliinae (extended, see Heppner 2003), Acrolepiinae (see Gaedike 1997) and Glyphipteriginae.
27. The order of superfamilies in Apoditrysia is very tentative; and their relationships are still unsettled.
28. Douglasiidae have been removed from Gracillarioidea, as two separate molecular analyses consistently placed them in Apoditrysia (Mutanen et
al. 2010; Kawahara et al. 2011).
29. The position of Gelechioidea in Apoditrysia follows Kaila (2004) and Mutanen et al. (2010). However, its sister group is completely unknown.
Therefore, we tentatively place this superfamily at the base of Apoditrysia. The present division into families is highly tentative; new molecular
results indicate the need for future changes.
30. As previously noted (Hodges 1998), there are senior names for Gelechiidae and Depressariinae that should be suppressed due to their infrequent
and forgotten use. At least at superfamily level, ICZN Article 35.5 probably applies. An application for suppression of these names will be
prepared by some of the present authors.
31. In addition to the groups included by Hodges (1998), Autostichidae now also contains Glyphidocerinae and Deocloninae. The monotypic genus
Oecia Walsingham, 1897, formerly in Schistonoeidae: Oeciinae, is here placed in subfamily Holcopogoninae (Gozmány 2000).
32. Elevation to family level and placement of Blastobasidae is updated from Baldizzone et al. (2006) and supported by Kaila (2004) and Bucheli &
Wenzel (2005).
33. Contains the subfamilies Oecophorinae and Deuterogoniinae (see Saito 2005). Stathmopodidae are here treated as a separate family.
34. Lypusidae were synonymised with Amphisbatidae Spuler, 1910 by Heikkilä & Kaila (2010), but Amphisbatidae is the junior synonym. Most of
the genera previously included in Amphisbatidae (e.g. Hodges 1998; Lvovsky 2011) have now been moved to Depressariinae in Elachistidae.
35. New synonym: Carcinidae Meyrick, 1906 (based on Carcina Hübner, 1825), invalid as it is a junior homonym of Carcinidae MacLeay, 1838
(based on Carcinus Leach, 1814) in Malacostraca: Decapoda, Brachyura.

214 · Zootaxa 3148 © 2011 Magnolia Press VAN NIEUKERKEN ET AL.: ORDER LEPIDOPTERA
 Family Elachistidae Bruand, 1850 (161 genera, 3,201 species) 36
Family Syringopaidae Hodges, 1998 (1 genus, 1 species)
Family Coelopoetidae Hodges, 1978 (1 genus, 3 species)
Family Stathmopodidae Janse, 1917 (44 genera, 408 species)
Family Epimarptidae Meyrick, 1914 (1 genus, 4 species) 37
Family Batrachedridae Heinemann & Wocke, 1876 (10 genera, 99 species)
Family Coleophoridae Bruand, 1850 (5 genera, 1,386 species) 38
Family Momphidae Herrich-Schäffer, 1857 (6 genera, 115 species)
Family Pterolonchidae Meyrick, 1918 (2 genera, 8 species)
Family Scythrididae Rebel, 1901 (30 genera, 669 species) 39
Family Cosmopterigidae Heinemann & Wocke, 1876 (135 genera, 1,792 species) 40
Family Gelechiidae Stainton, 1854 (500 genera, 4,700 species) 41
Superfamily Alucitoidea Leach, 1815 (2 families)
Family Tineodidae Meyrick, 1885 (12 genera, 19 species) 42
Family Alucitidae Leach, 1815 (9 genera, 216 species) 43
Superfamily Pterophoroidea Latreille, 1802 (1 family)
Family Pterophoridae Latreille, 1802 (90 genera, 1,318 species) 44
Superfamily Carposinoidea Walsingham, 1897 (2 families) 45
Family Copromorphidae Meyrick, 1905 (9 genera, 43 species)
Family Carposinidae Walsingham, 1897 (19 genera, 283 species)
Superfamily Schreckensteinioidea Fletcher, 1929 (1 family)
Family Schreckensteiniidae Fletcher, 1929 (2 genera, 8 species)
Superfamily Epermenioidea Spuler, 1910 (1 family)
Family Epermeniidae Spuler, 1910 (10 genera, 126 species) 46
Superfamily Urodoidea Kyrki, 1988 (1 family)
Family Urodidae Kyrki, 1988 (3 genera, 66 species)
Superfamily Immoidea Common, 1979 (1 family)
Family Immidae Common, 1979 (6 genera, 245 species)
Superfamily Choreutoidea Stainton, 1858 (1 family)
Family Choreutidae Stainton, 1858 (18 genera, 406 species)
Superfamily Galacticoidea Minet, 1986 (1 family)
Family Galacticidae Minet, 1986 (3 genera, 19 species)
Superfamily Tortricoidea Latreille, 1802 (1 family)
Family Tortricidae Latreille, 1802 (1,071 genera, 10,387 species) 47
Superfamily Cossoidea Leach, 1815 (7 families) 48

36. Elachistidae here include subfamilies Elachistinae, Depressariinae, Hypertrophinae, Ethmiinae, Aeolanthinae, Stenomatinae, Agonoxeninae and
Parametriotinae (Hodges 1998; Kaila 2004).
37. Previously considered a subfamily of Batrachedridae.
38. Updated from Baldizzone et al. (2006). In contrast to Hodges (1998), Coleophoridae, Momphidae, Blastobasidae and Pterolonchidae are here
treated as separate families.
39. Updated from Passerin d'Entreves & Roggero (2007).
40. Updated from Sinev (2002).
41. Based on an estimate for the largest subfamily Gelechiinae and actual counts of Physoptilinae, Dichomeridinae and Pexicopiinae.
42. Updated from Gielis (2003). Tineodidae appeared as paraphyletic in a molecular analysis (Mutanen et al. 2010). Based on that work,
Isonomeutis Meyrick, 1988 from New Zealand is moved here from Copromorphidae.
43. Updated from Gielis (2003).
44. Updated from Gielis (2003).
45. The name Copromorphoidea Meyrick, 1905, introduced by Meyrick (1928), is a junior synonym of Carposinoidea, which although introduced as
a superfamily by Diakonoff (1961), and several times used before 2000, is attributable to Walsingham (1897) as the author of the family on
which the name is based.
46. Updated from Gaedike (1996).
47. See Baixeras et al. (2010).
48. Cossoidea and Sesioidea were never monophyletic and were much intermixed in recent molecular analyses (Regier et al. 2009; Mutanen et al.
2010). Although the monophyly of these superfamilies together is also uncertain, we tentatively synonymise Sesioidea with Cossoidea. It is
possible that only by combining them with Zygaenoidea will a well-supported monophyletic group be eventually achieved.

ANIMAL BIODIVERSITY Zootaxa 3148 © 2011 Magnolia Press · 215

 Family Brachodidae Agenjo, 1966 (14 genera, 137 species) 49
Family Cossidae Leach, 1815 (151 genera, 971 species) 50
Family Dudgeoneidae Berger, 1958 (6 genera, 57 species) 51
Family Metarbelidae Strand, 1909 (18 genera, 196 species) 52
Family Ratardidae Hampson, 1898 (3 genera, 10 species)
Family Castniidae Boisduval, 1828 (34 genera, 113 species)
Family Sesiidae Boisduval, 1828 (154 genera, 1,397 species) 53
Superfamily Zygaenoidea Latreille, 1809 (12 families)
Family Epipyropidae Dyar, 1903 (9 genera, 32 species)
Family Cyclotornidae Meyrick, 1912 (1 genus, 5 species)
Family Heterogynidae Rambur, 1866 (1 genus, 10 species)
Family Lacturidae Heppner, 1995 (8 genera, 120 species)
Family Phaudidae Kirby, 1892 (3 genera, 15 species) 54
Family Dalceridae Dyar, 1898 (11 genera, 80 species)
Family Limacodidae Duponchel, 1845 (301 genera, 1,672 species)
Family Megalopygidae Herrich-Schäffer, 1855 (23 genera, 232 species)
Family Aididae Schaus, 1906 (2 genera, 6 species)
Family Somabrachyidae Hampson, 1920 (4 genera, 8 species)
Family Himantopteridae Rogenhofer, 1884 (11 genera, 80 species) 55
Family Zygaenidae Latreille, 1809 (170 genera, 1,036 species)
Clade Obtectomera Minet, 1986 (12 superfamilies, all following)
Superfamily Whalleyanoidea Minet, 1991 (1 family)
Family Whalleyanidae Minet, 1991 (1 genus, 2 species)
Superfamily Thyridoidea Herrich-Schäffer, 1846 (1 family)
Family Thyrididae Herrich-Schäffer, 1846 (93 genera, 940 species)
Superfamily Hyblaeoidea Hampson, 1903 (1 family)
Family Hyblaeidae Hampson, 1903 (2 genera, 18 species)
Superfamily Calliduloidea Moore, 1877 (1 family)
Family Callidulidae Moore, 1877 (7 genera, 49 species)
Superfamily Papilionoidea Latreille, 1802 (7 families) 56
Family Papilionidae Latreille, 1802 (32 genera, 570 species, 3/4) 57
Family Hedylidae Guenée, 1858 (1 genus, 36 species)
Family Hesperiidae Latreille, 1809 (570 genera, 4,113 species, 2/2)
Family Pieridae Swainson, 1820 (91 genera, 1,164 species, 5/6)
Family Riodinidae Grote, 1895 (1827) (146 genera, 1,532 species, 3/4)
Family Lycaenidae Leach, 1815 (416 genera, 5,201 species, 1/1)
Family Nymphalidae Rafinesque, 1815 (559 genera, 6,152 species, 11/21)
Superfamily Pyraloidea Latreille, 1809 (2 families) 58
Family Pyralidae Latreille, 1809 (1,055 genera, 5,921 species)
Family Crambidae Latreille, 1810 (1,020 genera, 9,655 species)
Superfamily Mimallonoidea Burmeister, 1878 (1 family)
Family Mimallonidae Burmeister, 1878 (27 genera, 194 species)

49. This includes also Pseudocossinae Heppner, 1984 (see Minet 1991).
50. Cossidae in the sense of the Handbook (Edwards et al. 1998) and Davis et al. (2008) were not monophyletic in recent molecular analyses (Regier
et al. 2009; Mutanen et al. 2010). Here they include only Zeuzerinae, Hypoptinae and Cossinae, almost as the treatment in Schoorl (1990). For
diversity of Old World taxa see Yakovlev (2011).
51. Dudgeoneidae now also include Cossulinae, following Mutanen et al. (2010), a relationship already suggested by Edwards et al. (1998).
52. Afrotropical diversity from De Prins & De Prins (2011b), to which two Oriental genera and ca. 20 species are added.
53. Diversity follows Pühringer & Kallies (2011).
54. Family status of Phaudidae is supported by molecular analyses (Niehuis et al. 2006; Mutanen et al. 2010).
55. Anomoeotidae Hering, 1937 are here regarded as subfamily of Himantopteridae (Mutanen et al. 2010).
56. Current placement and composition of butterflies (including Hesperiidae and Hedylidae) is strongly supported by both large molecular studies
(Regier et al. 2009; Mutanen et al. 2010). Diversity numbers follow G. Lamas’ database, only partly published (Lamas 2008).
57. Chilasa Moore [1881] is included in Papilio Linnaeus, 1758 as a junior synonym, following the most recent phylogenetic studies (Zakharov et
al. 2004; Simonsen et al. 2011).
58. Diversity based on two databases (Beccaloni et al. 2005; Nuss et al. 2010).

216 · Zootaxa 3148 © 2011 Magnolia Press VAN NIEUKERKEN ET AL.: ORDER LEPIDOPTERA
 Clade Macroheterocera Chapman, 1893 (5 superfamilies) 59
Superfamily Drepanoidea Boisduval, 1828 (3 families) 60
Family Cimeliidae Chrétien, 1916 (2 genera, 6 species) 61
Family Doidae Donahue & Brown, 1987 (2 genera, 6 species) 62
Family Drepanidae Boisduval, 1828 (122 genera, 660 species)
Superfamily Lasiocampoidea Harris, 1841 (1 family)
Family Lasiocampidae Harris, 1841 (224 genera, 1,952 species)
Superfamily Bombycoidea Latreille, 1802 (10 families) 63
Family Apatelodidae Neumoegen & Dyar, 1894 (10 genera, 145 species) 64
Family Eupterotidae Swinhoe, 1892 (53 genera, 339 species)
Family Brahmaeidae Swinhoe, 1892 (7 genera, 65 species) 65
Family Phiditiidae Minet, 1994 (4 genera, 23 species) 66
Family Anthelidae Turner, 1904 (9 genera, 94 species) 67
Family Carthaeidae Common, 1966 (1 genus, 1 species)
Family Endromidae Boisduval, 1828 (12 genera, 59 species) 68
Family Bombycidae Latreille, 1802 (26 genera, 185 species)
Family Saturniidae Boisduval, 1837 (169 genera, 2,349 species)
Family Sphingidae Latreille, 1802 (206 genera, 1,463 species, 2/2)
Superfamily Geometroidea Leach, 1815 (4 families)
Family Epicopeiidae Swinhoe, 1892 (9 genera, 20 species)
Family Sematuridae Guenée,1858 (6 genera, 40 species)
Family Uraniidae Leach, 1815 (90 genera, 686 species)
Family Geometridae Leach, 1815 (2,002 genera, 23,002 species) 69
Superfamily Noctuoidea Latreille, 1809 (6 families) 70
Family Oenosandridae Miller, 1991 (4 genera, 8 species)
Family Notodontidae Stephens, 1829 (704 genera, 3,800 species)
Family Erebidae Leach, 1815 (1,760 genera, 24,569 species) 71
Family Euteliidae Grote, 1882 (29 genera, 520 species) 72
Family Nolidae Bruand, 1847 (186 genera, 1,738 species)
Family Noctuidae Latreille, 1809 (1,089 genera, 11,772 species) 73

59. We prefer to use the term Macroheterocera for the well-supported moth clade identified in recent large molecular studies (Regier et al. 2009;
Mutanen et al. 2010), to avoid confusion with the popular term macrolepidopterans, now freed for popular use for a polyphyletic assemblage of
larger moths and butterflies.
60. Inclusion of Cimeliidae and Doidae follows tentatively recent molecular results (Regier et al. 2009; Mutanen et al. 2010) and is supported by
some morphological characters.
61. See Yen & Minet (2007) for synonymy of Axiidae Rebel, 1919 (a homonym of Axiidae Huxley, [1879] in Malacostraca: Decapoda) and its
replacement by the name Cimeliidae.
62. Doidae are moved from Noctuoidea, see Drepanoidea above.
63. Classification of Bombycoidea follows Zwick et al. (2011).
64. Removed from Bombycidae and raised to family by Zwick (2008).
65. Lemoniidae were synonymized with Brahmaeidae by Zwick (2008).
66. Removed from Bombycidae and raised to family by Zwick et al. (2011).
67. Removed from Lasiocampoidea by Zwick et al. (2011).
68. Zwick et al. (2011) synonymized three groups with Endromidae: Mirinidae, Primostictini and Oberthueriini (previously placed in Bombycidae -
Prismostictinae).
69. Diversity updated from Scoble & Hausmann (2007).
70. The classification of Noctuoidea follows the recent studies of Zahiri et al. (2011) and Lafontaine & Schmidt (2010). Doidae are moved from
Noctuoidea to Drepanoidea (see there). The six families recognized were all highly supported in the molecular analyses (Zahiri et al. 2011).
Molecular analyses are as yet inconclusive in establishing robust phylogenetic relationships among the four families of quadrifid noctuoids, so
the traditional sequence based on morphology is followed, placing Euteliidae after Erebidae and Nolidae before Noctuidae.
71. Erebidae include the former families Arctiidae Leach, 1815 and Lymantriidae Hampson, 1893 as subfamilies. Micronoctuidae Fibiger, 2005 is
subordinate within Hypenodinae (Lafontaine & Schmidt 2010; Zahiri et al. 2011). Diversity is updated from Poole (1989), numbers for arctiines
and lymantriines are estimates.
72. Euteliidae comprise Euteliinae and Stictopterinae (Zahiri et al. 2011). Diversity is updated from Poole (1989).
73. Noctuidae comprise essentially the trifine noctuoids (see Lafontaine & Schmidt 2010; Zahiri et al. 2011). Diversity is updated from Poole
(1989).

ANIMAL BIODIVERSITY Zootaxa 3148 © 2011 Magnolia Press · 217

Acknowledgements

The authors for the Lepidoptera chapter are grateful to Dana Campbell (University of Maryland) for opening the Leptree
forum for discussions amongst us (Leptree Team 2011). Support to Joaquín Baixeras was provided by grant CGL2008-
00605 of the Spanish Government (Ministerio de Ciencia e Innovación). Jurate De Prins is grateful to the Belgian Sci-
ence Policy Office for granting her study on Gracillariidae (contract MO/37/022) and the Belgian Biodiversity Platform
(head Dr. André Heughebaert) for the kind technical assistance to make the taxonomic data available online. David Lees
was supported by INRA-EFPA during preparation of this work. We thank Thomas Sobczyk (Hoyerswerda, Germany) for
providing data from his in press catalogue.

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Author addresses

Erik J. van Nieukerken (co-ordination Lepidoptera, Nepticuloidea, Adeloidea, various smaller groups) Netherlands Centre for
Biodiversity Naturalis, PO Box 9517, NL-2300 RA Leiden Netherlands; [email protected] (corresponding author)
Lauri Kaila (Lepidoptera classification, Gelechioidea) Finnish Museum of Natural History, P.O. Box 17, FI-00014 University
of Helsinki Finland; [email protected]
Ian J. Kitching (co-ordination Lepidoptera, Bombycoidea, Lasiocampoidea, Noctuoidea) Department of Entomology, Natural
History Museum, Cromwell Road, London SW7 5BD United Kingdom; [email protected]
Niels P. Kristensen (Lepidoptera classification, non-Heteroneura) Natural History Museum of Denmark, University of Copen-
hagen, Universitetsparken 15, DK-2100 Copenhagen Ø Denmark; [email protected]
David C. Lees (Lepidoptera classification, Unassigned early lepidopterans, Micropterigidae, Callidulidae) Department of Ento-
mology, Natural History Museum, Cromwell Road, London SW7 5BD United Kingdom/UR 633 Zoologie Forestière,
Institut National de la Recherche Agronomique, Orléans 45075 France; [email protected]
Joël Minet (Lepidoptera classification, Unassigned early lepidopterans, Callidulidae, Geometroidea) Muséum National d'His-
toire Naturelle, Département Systématique et Evolution, UMR CNRS 7205, Case Postale 50 (Entomologie), FR-75231
Paris Cedex 05 France; [email protected]
Charles Mitter (Lepidoptera classification) Department of Entomology, University of Maryland, College Park MD 20742 USA;
[email protected]
Marko Mutanen (Lepidoptera classification) Zoological Museum, Department of Biology, University of Oulu, PO Box 3000,
FI-90014 Oulu Finland; [email protected]
Jerome C. Regier (Lepidoptera classification) Institute for Bioscience and Biotechnology Research & Department of Entomo-
logy, University of Maryland, 5140 Plant Sciences Bldg. College Park, MD 20742 USA; [email protected]
Thomas J. Simonsen (co-ordination Lepidoptera; Exoporia, Papilionidae, Nymphalidae, Pyraloidea) Department of Entomo-
logy, Natural History Museum, Cromwell Road, London SW7 5BD United Kingdom; [email protected]
Niklas Wahlberg (Lepidoptera classification) Laboratory of Genetics, Department of Biology, University of Turku, FI-20014
Turku Finland; [email protected]
Shen-Horn Yen (Lepidoptera: Zygaenoidea, Thyrididae, Crambidae) Department of Biological Sciences, National Sun Yat-Sen,
University, 70 Lien Hai Rd, Kaohsiung 804 Taiwan; [email protected]
Reza Zahiri (Lepidoptera: Noctuoidea) Laboratory of Genetics, Department of Biology, University of Turku, FI-20014 Turku
Finland; [email protected]
David Adamski (Lepidoptera: Blastobasidae, Autostichidae) Department of Entomology, National Museum of Natural History,
Smithsonian Institution Washington, DC 20013-7012 USA; [email protected]
Joaquin Baixeras (Lepidoptera: Tortricoidea) Cavanilles Institute of Biodiversity and Evolutionary Biology, University of
Valencia, P.O.Box 22085, ES- 46071 Valencia Spain; [email protected]
Daniel Bartsch (Lepidoptera: Sesiidae) State Museum of Natural History Stuttgart, Rosenstein 1, DE-70191 Stuttgart Germany;
[email protected]
Bengt Å. Bengtsson (Lepidoptera: Scythrididae) Lokegatan 3, SE-386 93 Färjestaden Sweden; [email protected]
John W. Brown (Lepidoptera: Tortricoidea) SEL, USDA, Smithsonian Institution, P.O. Box 37012, National Museum Natural
History, E-517, MRC 168, Washington, DC 20013-7012 USA; [email protected]
Sibyl Rae Bucheli (Lepidoptera: Gelechioidea) Department of Biological Sciences, Sam Houston State University, Box 2116,
Huntsville TX 77341 USA; [email protected]
Donald R. Davis (Lepidoptera: Neopseustidae, Tineoidea, Epipyropidae) Entomology, NHB 105, P.O. Box 37012, Smithso-
nian Institution, Washington, DC, USA, 20013-7012; [email protected]
Jurate De Prins (Lepidoptera: Gracillariidae) Royal Museum for Central Africa, Leuvensesteenweg 13, BE-3080 Tervuren Bel-
gium; [email protected]
Willy De Prins (Lepidoptera: Gracillariidae) Netherlands Centre for Biodiversity Naturalis, PO Box 9517, 2300 RA Leiden
The Netherlands; [email protected]
Marc E. Epstein (Lepidoptera: Zygaenoidea) California Dept. of Food & Agriculture, Plant Pest Diagnostics Branch, 3294
Meadowview Ave, Sacramento, CA 95832-1448 USA; [email protected]
Patricia Gentili-Poole (Lepidoptera: Cossoidea) Department of Entomology, Smithsonian Institution, Washington DC 20013-

220 · Zootaxa 3148 © 2011 Magnolia Press VAN NIEUKERKEN ET AL.: ORDER LEPIDOPTERA
 7012 USA; [email protected]
Cees Gielis (Lepidoptera: Pterophoridae, Alucitoidea) Netherlands Centre for Biodiversity Naturalis, PO Box 9517, NL-2300
RA Leiden Netherlands; [email protected]
Peter Hättenschwiler (Lepidoptera: Psychidae) Seeblickstrasse 4, CH-8610 Uster Switzerland;
[email protected]
Axel Hausmann (Lepidoptera: Geometroidea) Bavarian State Collection of Zoology, Münchhausenstr. 21, DE-81247 Munich
Germany; [email protected]
Jeremy D. Holloway (Lepidoptera: Noctuoidea, various Apoditrysia) Department of Entomology, Natural History Museum,
Cromwell Road, London SW7 5BD United Kingdom; [email protected]
Axel Kallies (Lepidoptera: Brachodidae, Sesiidae) The Walter and Eliza Hall Institute of Medical Research, 1G Royal Parade,
3050 Parkville Victoria Australia; [email protected]
Ole Karsholt (Lepidoptera: Gelechiidae) Natural History Museum of Denmark, University of Copenhagen, Universitetsparken
15, DK-2100 Copenhagen Ø Denmark, [email protected]
Akito Y. Kawahara (Lepidoptera: Gracillarioidea) McGuire Center For Lepidoptera and Biodiversity, Florida Museum of Nat-
ural History, University of Florida, SW 34th Street and Hull Road, PO Box 112710, Gainesville, FL 32611-2710 USA;
[email protected]
Sjaak (J.C.) Koster (Lepidoptera: Gelechioidea) Netherlands Centre for Biodiversity Naturalis, PO Box 9517, NL-2300 RA
Leiden Netherlands; [email protected]
Mikhail V. Kozlov (Lepidoptera: Adelidae) Section of Ecology, University of Turku, FI-20014 Turku, Finland; [email protected]
J. Donald Lafontaine (Lepidoptera: Noctuoidea) Canadian National Collection of Insects, Arachnids, and Nematodes , Agricul-
ture and Agri-Food Canada, 960 Carling Avenue, Ottawa ON K1A 0C6 Canada; [email protected]
Gerardo Lamas (Lepidoptera: Papilionoidea) Museo de Historia Natural, Universidad Nacional Mayor de San Marcos, Apar-
tado 14-0434, Lima-14 Peru; [email protected]
Jean-François Landry (Lepidoptera: Gelechioidea) Canadian National Collection of Insects, Arachnids, and Nematodes , Agri-
culture and Agri-Food Canada, 960 Carling Avenue, Ottawa ON K1A 0C6 Canada; [email protected]
Sangmi Lee (Lepidoptera: Gelechiidae) Mississippi Entomological Museum, PO Box 9775, Mississippi State University, Mis-
sissippi State, MS 39762 USA; [email protected]
Matthias Nuss (Lepidoptera: Pyraloidea) Senckenberg Naturhistorische Sammlungen Dresden, Museum für Tierkunde,
Königsbrücker Landstr. 159, DE-01109 Dresden Germany; [email protected]
Kyu-Tek Park (Lepidoptera: Lecithoceridae) McGuire Center for Lepidoptera and Biodiversity, Florida Museum of Natural
History, University of Florida, Gainesville, FL 32611-2710 USA; [email protected]
Carla Penz (Lepidoptera: Nymphalidae) Department of Biological Sciences, University of New Orleans, 2000 Lakeshore
Drive, New Orleans, LA 70148 USA; [email protected]
Jadranka Rota (Lepidoptera: Choreutidae, Millieriidae) Natural History Museum of Denmark, University of Copenhagen, Uni-
versitetsparken 15, DK-2100 Copenhagen Ø Denmark; [email protected]
Alexander Schintlmeister (Lepidoptera: Notodontidae) Calberlastr. 3, DE-01326 Dresden Germany; [email protected]
B. Christian Schmidt (Lepidoptera: Noctuoidea) Canadian Food Inspection Agency, Canadian National Collection of Insects,
Arachnids and Nematodes, 960 Carling Avenue, Ottawa, ON, Canada K1A 0C6; [email protected]
Jae-Cheon Sohn (Lepidoptera: Yponomeutoidea) Department of Entomology, University of Maryland, 4112 Plant Sciences
Bldg., College Park, MD 20742 USA; [email protected]
M. Alma Solis (Lepidoptera: Pyraloidea) SEL, USDA, Smithsonian Institution, P.O. Box 37012, National Museum Natural
History, E-517, MRC 168, Washington, DC 20013-7012 USA; [email protected]
Gerhard M. Tarmann (Lepidoptera: Zygaenidae) Tiroler Landesmuseum Ferdinandeum, Naturwissenschaftliche Sammlungen,
Feldstrasse 11a, AT-6020 Innsbruck Austria; [email protected]
Andrew D. Warren (Lepidoptera: Hesperiidae) McGuire Center for Lepidoptera and Biodiversity, Florida Museum of Natural
History, University of Florida, SW 34th Street and Hull Road, P. O. Box 112710, Gainesville, FL 32611-2710;
[email protected]
Susan Weller (Lepidoptera: Noctuoidea) J.F. Bell Museum of Natural History, 10 Church St, S.E., Room 300, University of
Minnesota, St. Paul MN 55455 USA; [email protected]
Roman V. Yakovlev (Lepidoptera: Cossoidea) South-Siberian Botanical Garden, Altai State Uni., ul. Chkalova 57-81, RU-
656049 Barnaul Russia; [email protected]
Vadim V. Zolotuhin (Lepidoptera: Lasiocampoidea, Bombycoidea) Ablukova 13-12, RU- 432005 Uljanovsk Russia
[email protected]
Andreas Zwick (Lepidoptera: Lasiocampoidea, Bombycoidea) State Museum of Natural History Stuttgart, Rosenstein 1, DE-
70191 Stuttgart Germany; [email protected]

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