Monophyly of Noctuoidea
Noctuoidea is currently recognized as a large superfamily comprising some 70,000 described species (Kitching & Rawlins 1998). Monophyly of Noctuoidea, based on the presence of a single apomorphic character, the metathoracic tympanal organ (Miller 1991) and its associated abdominal structures (Kitching & Rawlins 1998), seems well established (Nielsen 1989; Common 1990; Miller 1991; Scoble 1992; Kitching & Rawlins 1998; Mitchell et al. 2006), but in general the limits and content of the constituent families, and the relationships among and within these, is poorly understood (Mitchell et al. 2006). Furthermore, the presence of two MD setae on the larval metathorax is a consistent and reliable synapomorphy of all Noctuoidea except the Australian family, Oenosandridae (Kitching & Rawlins 1998).
Moths of this superfamily constitute the most speciose and cosmopolitan lineage of all Lepidoptera, with adults and larvae exhibiting an often bewildering diversity of sizes (from very small to very large), coloration, behaviour and ecology (Kitching & Rawlins 1998). Chaetosemata are always absent. Wing coupling is effected by a frenulo-retinacular system (Scoble 1992). The eggs of Noctuoidea are of the upright variety. The larvae are cylindrical with prolegs bearing crochets arranged in a mesoseries.
Fossil records of noctuoid lineages
Grimaldi & Engel (2005) recorded the earliest fossil of Noctuoidea is an unnamed specimen from the Tertiary (Paleocene), about 55 Mya, of Denmark (Rust 2000). The other one is Stauropolia nekrutenkoi probably an arctiine is from the Tertiary (Miocene) of the Caucasus, about 15 Mya (Skalski 1988). Kitching & Rawlins (1998) commented that the fossil record of Noctuoidea is too meagre and young (Middle to Late Tertiary) to be phylogenetically informative. A few genera of Holarctic fossils of uncertain affinity have been proposed: Arctiites, Stauropolia and Oligamatites (Noctuidae, Arctiinae); Cerurites (Notodontidae); Noctuites and Phylledestes (Noctuidae).
An egg from the Cretaceous has been attributed to Noctuidae (Gall & Tiffney 1983), but the determination is not based on a diagnostic apomorphy for noctuoid eggs and remains dubious (Kitching & Rawlins 1998). This fossil moth egg was found in 75-million-year-old sediments in Massachusetts. The egg is positively assigned to the moth family Noctuidae and extends the fossil record of this family back into the Cretaceous. It turns out that Noctuidae family moths have special organs for detecting the ultrasonic cries of insect-hunting bats. The fossil record of the bats, however, only goes back to the early Eocene, perhaps 20 million years after the Noctuidae moths. Since no other insect predators like bats existed, it would seem that the moths developed these special organs in anticipation of the bats!
Very recent unpublished investigations have revealed that some groups of Papilionoidea may be as old as 85 Mya, which more or less coincides with the major angiosperm radiation. Assuming the origin of Noctuoidea is more ancient than that of butterflies (an hypothesis that remains to be severely tested), it is reasonable to suppose that noctuoids radiated before 85 Mya.
Trifid and Quadrifid Noctuoidea
It should be noted that within Noctuoidea, there are two basic patterns of forewing venation. In the plesiomorphic state, MA2 (=M2) arises midway between MA1 (=M1) and MP1 (=M3), giving an apparent trifurcation at the lower corner of the discal cell (Fig. Trifid Noctuoid, Notodontidae, Thaumetopoeinae). This is referred to as the trifid condition and occurs in Oenosandridae, Doidae and Notodontidae. In contrast, the form in which MA2 arises very close to, connate with or stalked with MP1, giving an apparent four-branched pattern, is referred to as the quadrifid condition (Fig. Quadrifid Noctuoid, Arctiidae: Arctiinae) and occurs in all other noctuoids, except some derived lineages showing fewer branches through fusion.
Noctuoidea classification
Numerous classifications for the family groups of Noctuoidea have been proposed. The fundamental distinction between the different systems is based on the use of different (occasionally plesiomorphic) characters in phylogenetic reconstruction. For example, Miller (1991), proposed seven families: Oenosandridae, Doidae, Notodontidae, Lymantriidae, Arctiidae, Aganaidae and Noctuidae. Scoble (1992) considered six families, comprising Oenosandridae, Doidae, Notodontidae, Lymantriidae, Arctiidae and Noctuidae (including Aganainae). One of the most comprehensive studies of Noctuoidea by Kitching & Rawlins (1998) recognized three fundamental lineages of Noctuoidea: Oenosandridae, Doidae + Notodontidae, and the quadrifid families (they treated the following families: Arctiidae, Lymantriidae, Noctuidae, Nolidae and Pantheidae). Miller (1991) favoured a clade comprising Oenosandra and Doidae + Notodontidae, on the basis of two synapomorphies: presence of a stipital lobe in the larvae and presence of socii, although Kitching and Rawlins (1998) stated both features are problematic.
Grimaldi & Engel (2005) followed Miller (1991) and stated that the superfamily includes two small families, the Oenosandridae (with three Australian species that feed on Myrtaceae), and the Neotropical Doidae, which feed on Euphorbiaceae and are closely related to the Notodontidae, and four large families: Notodontidae, Lymantriidae, Arctiidae and Noctuidae.
The higher classification of the superfamily was recently and comprehensively reviewed by Fibiger and Lafontaine (2005). They proposed a new classification for the superfamily that included 10 families: Oenosandridae, Doidae, Notodontidae, Strepsimanidae, Nolidae, Lymantriidae, Arctiidae, Erebidae, Micronoctuidae, and Noctuidae. Fibiger & Lafontaine (2005) also presented a cladogram depicting the possible relationships of families of Noctuoidea (Cladogram Noctuoidea: Fibiger & Lafontaine 2005).
Most recently, Lafontaine & Fibiger again proposed another classification system for the families of the Noctuoidea with a quadrifid forewing venation (Nolidae, Strepsimanidae, Arctiidae, Lymantriidae, Erebidae, and Noctuidae), mainly based on recent classifications and the distribution of derived character states. In this new system, Nolidae, Strepsimanidae, Arctiidae, Lymantriidae, and Erebidae sensu Fibiger & Lafontaine (2005) were downgraded to the subfamily status within an expanded family concept of Noctuidae. Consequently, the superfamily Noctuoidea now comprises only the families Oenosandridae, Doidae, Notodontidae, Micronoctuidae and Noctuidae.
Future of Noctuoidea classification
Prominent Noctuidae specialists made decision at a workshop on Noctuidae classification at the Societas Europaea Lepidopterologica (SEL) Congress at Korsør, Denmark, in June, 2002, to provide a review of family-group names in order to elucidate nomenclatural decisions, which were eventually published by Kühne & Speidel (2004) and Speidel & Naumann (2005). Speidel & Naumann (2005) concluded their paper with a number of important points of view concerning the future of Noctuoidea systematics, and emphasized that there is no difference between so-called larval and imaginal systematics, even though this impression could easily be gained from the study of some papers by Beck (e.g. Beck 1989). Systematics has always tried to incorporate the study of all life stages. However, there is an evident contradiction between phylogenetic and typological systematics. Polyphyletic and paraphyletic groups must be broken up to obtain a phylogenetic system. Furthermore, in addition to larval and adult characters, there is now another important new set of characters provided by molecular sequences, the utility of which is still difficult to determine in detail. At present, and despite all the phylogenetic work of the last decade, we are still far from recognizing the major phylogenetic lines within Noctuoidea.
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