FNZ 14 - Lepidoptera - annotated catalogue and keys to family group taxa - Classification
Dugdale, JS 1988. Lepidoptera - annotated catalogue, and keys to family-group taxa. Fauna of New Zealand 14, 264 pages.
(
ISSN 0111-5383 (print),
;
no.
14.
ISBN 0-477-02518-8 (print),
).
Published 23 Sep 1988
ZooBank: http://zoobank.org/References/B69FC31C-8562-4047-AD44-3A37E38FA872
Classification
Relationships of Lepidoptera
The order Lepidoptera, with between 100,000 and 200,000 known species, is currently regarded as a member of the "Panorpoid complex" or Mecopteroidea, which comprises the superorders Antliophora (Mecoptera + Siphonaptera + Diptera) and Amphiesmenoptera (Trichoptera + Lepidoptera) (see Kristensen 1984, p. 145). Lepidoptera are distinguished from their sister-group Trichoptera by the following apomorphies or unique character states:
- vestiture of wings and body largely made up of broad, overlapping scales;
- head lacking a median ocellus;
- apical segment of labial palpus with a group of receptors in a depression (vom Rath's organ).
Some other structures, such as the fore-tibial epiphysis, are not always present; even vom Rath's organ may not be in a pit, as in Southern Hemisphere Roeslerstammiidae (I.F.B. Common, pers. comm.). There is, in fact, only a very small suite of character states that distinguish all Lepidoptera from all Trichoptera, and Kristensen (1984, p. 156) has pointed out the similarity in habitus between less advanced Trichoptera and the lepidopteran Agathiphaga Dumbleton.
Higher classification within Lepidoptera
The higher classification within this order is currently undergoing extensive re-examination, following on from the last intensive summary (Common 1975). The situation is rendered difficult because of two factors:
- around 98% of known Lepidoptera belong to one division (Ditrysia), of relatively uniform structural organisation;
- the remaining 1 - 2% show a great variety of structural and genital organisation, with often profound differences between groups.
There is a consensus (Kristensen 1984, Nielsen 1985, Davis 1986) that Lepidoptera phylogeny is an example of Hennig's "additive typogenesis" (Nielsen 1985, p. 1 and fig. 1). This implies a gradual acquisition of group characters from Zeugloptera onwards, culminating in the endoporian ditrysian suite of characters. However, "this relatively simple picture has become modified and much more complicated by the considerable amount of new information ... available during the last two decades" (Nielsen 1985, p. 2).
The following group-names have been used above the superfamily level (see Kristensen 1984, p. 186, fig. 13): Zeugloptera (includes Micropterigoidea); Aglossata (includes Agathiphagoidea); Heterobathmiina (includes Heterobathmioidea); Glossata, divided into Dacnonypha (haustellum without intrinsic musculature) and Myoglossata (with intrinsic musculature); the Myoglossata including the Neopseustina (Neopseustoidea); and the Neolepidoptera, including the Exoporia (Mnesarchaeoidea, Hepialoidea) and the Heteroneura (Nepticuloidea, Incurvarioidea, Palaephatoidea); the Eulepidoptera including the Etimonotrysia (Tischerioidea); and Ditrysia (all other superfamilies). Note that Glossata, Myoglossata, Neolepidoptera, and Eulepidoptera (Minet 1984, p. 147) are clades. Opinion is divided (cf. Minet 1984, Nielsen 1982, 1985, and Davis 1986) over the monophyly or otherwise of such categories as Heteroneura and Neolepidoptera, and, for that matter, of the position of the taxa Heterobathmiina and Exoporia (left hanging by Minet 1984, fig. 26), and whether the Zeugloptera alone, or Zeugloptera + Aglossata are the sister-group of all other Lepidoptera (see discussion by Kristensen 1984, p. 166).
Comprehensive discussions of Heteroneura phylogeny are given by Common (1975), Davis (1986, pp. 55 - 61), and Nielsen (1982, 1985a, 1985b). By and large the argument involves two assumptions: (a) that changes in frenular structures arose once; and (b) that the internal change from a ventral to a dorsal common oviduct in relation to the copulatory chamber (Dugdale 1974) arose independently in Exoporia and Ditrysia (Common 1975).
The classificatory outline given below is largely derived from Nielsen (1985a, pp. 15 - 16; 1985b, p. 142), Minet (1986), and Nielsen & Common (in prep.), but no categories above superfamily are listed except where the higher category includes only one superfamily. In such instances the higher term is given only as a recognition term, of possible use to the reader in literature searches.
Within the division (Davis 1986) or phalanx (Minet 1984, 1986) Ditrysia some 28 superfamilies are recognised (Minet 1983; Nielsen & Common, in prep.). Brock (1971) noted two groupings of superfamilies; these were refined by Kyrki (1983) and Minet (1983), largely on the basis of the structure of the thoracic / abdominal junction in adult moths. The term Apoditrysia (Minet 1983, p. 201) includes all ditrysian superfamilies with a tortricoid basisternite, and excludes those with a tineoid basisternite, i.e., Tineoidea, Yponomeutoidea, and Gelechioidea. As a concept, it still requires testing over a greater range of species, and its relation to metafurcasternal structures (Davis 1986) and prothoracic structures (Minet 1984) needs more precise evaluation.
Minet (1986) gives a revised outline of Lepidoptera classification, and reduces the number of ditrysian superfamilies to 26. His removal of Choreutidae from Sesioidea - as "Ditrysia à affinites inconnue ou uncertaine" (p. 292, and note 61, p. 306) - is of relevance to the New Zealand fauna. Minet's outline does not include Hedyloidea, newly recognised as a group related to Papilionoidea (Scoble 1986).
Superfamilies of Lepidoptera, and their families
Those superfamilies and families absent from New Zealand have a minus sign following them; where the sign is in parentheses, the family is represented by introduced species only; e.g., Cossidae (-).
- Micropterigoidea: Micropterigidae [Zeugloptera]
- Agathiphagoidea: Agathiphagidae - [Aglossata]
- Heterobathmioidea: Heterobathmiidae -
- Eriocranioidea: Eriocraniidae -, Lophocoronidae -, Acanthopteroctetidae - [Dacnonypha]
- Neopseustoidea: Neopseustidae -
- Nepticuloidea: Nepticulidae; Opostegidae -
- Mnesarchaeoidea: Mnesarchaeidae [Exoporia]
- Hepialoidea: Hepialidae in the strict sense; "primitive Hepialidae" -, Anomosetidae -, Neotheoridae -, Palaeosetidae -, Prototheoridae - [Exoporia]
- Incurvarioidea: Prodoxidae [?]; Adelidae -, Cecidoseidae -, Crinopterigidae -, Incurvariidae -, Heliozelidae -
- Palaephatoidea: Palaephatidae -
- Tischerioidea: Tischeriidae -
- [Ditrysia: "non-apoditrysian" superfamilies]
- Tineoidea: Tineidae, Psychidae, Arrhenophanidae -, Eriocottidae -, Pseudarbelidae -, Roeslerstammiidae, Gracillariidae, Bucculatrigidae -, Douglasiidae -
- Yponomeutoidea: Yponomeutidae (including Ypsolophidae, Plutellidae in the sense of Nielsen & Common, in prep.), Glyphipterigidae, Lyonetiidae (including Bedelliinae), Heliodinidae -
- Gelechioidea: Gelechiidae, Coleophoridae, Batrachedridae (in the sense of Nielsen & Common, in prep.), Cosmopterigidae, Blastobasidae (including Symmocidae, in the sense of Hodges 1978, p. 7) (-), Momphidae, Oecophoridae (Depressariinae, Oecophorinae, Stathmopodinae, Stenomatinae, Xyloryctinae, Peleopodinae in the sense of Hodges 1974 (-)), Lecithoceridae, Scythrididae, Holcopogonidae -, Elachistidae, Agonoxenidae -
- [Ditrysia: Apoditrysia]
- Cossoidea: Cossidae (-), Metarbelidae -, Dudgeoneidae -, Ratardidae -
- Castnioidea: Castniidae - (Minet 1986 includes Castniidae in Sesioidea)
- Sesioidea: Sesiidae (-), Choreutidae, Brachodidae -
- Tortricoidea: Tortricidae (Tortricinae, Chlidanotinae, Olethreutinae)
- Zygaenoidea: Zygaenidae -, Heterogynidae -, Phaudidae -, Megalopygidae -, Chrysopolomidae -, Somabrachyidae -, Limacodidae -, Dalceridae -, Epipyropidae -, Cyclotornidae -
- Immoidea: Immidae -
- Copromorphoidea: Copromorphidae, Carposinidae
- Epermenioidea: Epermeniidae
- Schreckensteinioidea: Schreckensteiniidae -
- Alucitioidea: Tineodidae -, Alucitidae -
- Pterophoroidea: Pterophoridae
- Hyblaeoidea: Hyblaeidae -
- Hedyloidea: Hedylidae -.
- Hesperoidea: Hesperiidae -
- Papilionoidea: Nymphalidae, Lycaenidae, Papilionidae -, Pieridae (-)
- Bombycoidea: Bombycidae (-), Saturniidae (-), Anthelidae -, Apatelodidae -, Endromidae -, Eupterotidae -, Hibrildidae -, Lacosomidae -, Lasiocampidae -, Lemoniidae -, Brahmaeidae -, Carthaeidae -, Mirinidae, Oxytenidae -, Cercophanidae -
- 'Sphingoidea'. Sphingidae (now regarded as bombycoids by Minet 1986)
- Thyridoidea: Thyrididae
- Calliduloidea: Callidulidae -, Pterothysanidae -
- Pyraloidea: Pyralidae (Galleriinae, Phycitinae, Pyralinae), Crambidae (Crambinae, Pyraustinae, Nymphulinae, Scopariinae, Musotiminae)
- Geometroidea: Geometridae (Ennominae, Geometrinae -, Larentiinae, Oenochrominae, Sterrhinae)
- Axioidea: Axiidae -
- Drepanoidea: Drepanidae (Thyatirinae) (-)
- Uranioidea: Uraniidae (including Epipleminae) -
- Mimallonoidea: Mimallonidae -
- Noctuoidea: Arctiidae, Ctenuchidae, Noctuidae (Agaristinae, Acronictinae, Cuculliinae, "Catocalinae" (-), Chloephorinae (-), Euteliinae -, Hadeninae, Heliothinae (-), Herminiinae -, "Hypeninae", Hypenodinae, Noctuinae, Nolinae, "Ophiderinae", Plusiinae (-), Stictopterinae -), Notodontidae -, Thaumetopoeidae -, Dioptidae -, Thyretidae -, Lymantriidae (-), Aganaidae -
- (Families of unknown position: Sematuridae -, Apoprogonidae -, Galacticidae -, Pterolonchidae -, Lathrotelidae -, Epicopeidae -)
- Superfamily unknown: Titanomis Meyrick; "Lysiphragma" argentaria Salmon (both apoditrysian; see p. 214).
- Families in which revisionary work has been or is being done in a world context: Micropterigidae, Nepticulidae, Mnesarchaeidae, Hepialidae*, Tineidae*, Tortricidae*, Gelechiidae (part), Crambidae (Crambinae*).
- Families in which some revisionary work has been or is being done in a local context: Noctuidae*, Geometridae*, Yponomeutidae (in a broad sense)*, Choreutidae, Momphidae, Tineidae*, Psychidae, Elachistidae, Arctiidae, Nymphalidae.
- Families newly recognised in New Zealand: Epermeniidae, Roeslerstammiidae, ?Prodoxidae.
- Families most in need of revision once overseas classification makes this practicable (usually families with a confused or contradictory subfamily classification or little recent overseas information): Oecophoridae* (including Depressariidae*), Noctuidae*, and the taxa Titanomis and "Lysiphragma" argentaria.
- Families that practicably can be revised now: Psychidae*, Tineidae*, Gracillariidae*, Yponomeutidae*, Glyphipterigidae*, Crambidae (Scopariinae*), Gelechiidae*, Carposinidae*, Elachistidae, Cosmopterigidae*, Choreutidae, Pyralidae*, Geometridae*.