Importance of Noctuidae
The family Noctuidae, sometimes known as owlets, is the most speciose radiaition of Lepidoptera, containing more than 35,000 described species in some 4,200 described genera (Kitching & Rawlins 1998) and numerous undescribed taxa, particularly from tropical areas. Furthermore, the larvae of many noctuid genera (e.g. Spodoptera, Heliothis, Euxoa, Earias and Trichoplusia), well known as army-worms, cutworms, bollworms and stem borers, are extremely important economically and cause many millions of Euros worth of crop losses each year. At present, their control, still mainly through the use of chemicals, is also expensive. A predictive classification is thus an essential prerequisite to the efficient use of the resources available to combat the pest species and also to minimize environmental contamination.
Noctuidae is monophyletic or paraphyletic group?
Most of the subfamilies in the classification of Hampson (1903- 1913) are now recognized as unnatural (Beck 1960, 1991, 1992; Kitching 1984; Lafontaine & Poole 1991; Speidel & Naumann 1996; Kitching & Rawlins 1998; Fibiger & Lafontaine 2005; Mitchell et al. 2006). Indeed, the composition and monophyly of most is still open to question, as is their number. The morphology of the immature stages, as well as the adults, is astonishingly poorly known and yet such information will be critical to a defensible noctuid phylogeny (Kitching & Rawlins 1998).
Yela & Kitching (1999) reviewed Noctuidae phylogeny based on morphological characterisation at the subfamily level, citing previous authors' concerns regarding the difficulties of noctuid phylogeny. For example, one of the most intractable problems in Lepidoptera classification has been the phylogeny of the quadrifid lineages of Noctuoidea, a group that has been notoriously difficult to characterise (both from Kitching & Rawlins 1998). Kitching (1984), in the publication that represented the inflexion point towards a cladistic approach to studies on noctuid phylogeny, also cited similar sentences. For example, “It is exceptional to find any two authors who use the same combination of subfamily names within the Noctuidae (Nye 1975)”; and “The classification [of the family Noctuidae] rests in a state of great confusion, and few authors appear to hold similar views regarding the suprageneric taxonomy. I have concluded after spending much time (perhaps I should say wasting time) on the problem, that it is impossible to present ...a correct suprageneric classification (Zimmerman 1958)”. Moreover, Poole (1995) added: “... the family is so large and the interpretation of characters so difficult that no one has tried to construct a natural classification reflecting the phylogeny of the family”. Finally, Speidel et al. (1996) stated that “ ... our understanding of the phylogenetic affinities of the Noctuidae to other noctuoid families, as well as interrelations among the various noctuid subfamilies, is still in its infancy”. Hence Yela & Kitching's (1999) comment that: “Thus, it can be easily understood that a review is not a simple task”.
Furthermore, it should be noted that the current state of the systematics of Noctuidae, which remains too Holarctic-oriented, is also due to the vastness of the group. Unfortunately, very little information is available on Neotropical noctuids (in particular quadrifines) because much of the basic alpha-taxonomy is still unclear. There are genera that include tens of described species but one will not ever find even a single illustration of one of those species to serve as a starting point for revision. It has been acknowledged by Fibiger & Lafontaine (2005) that a fuller study of the southern hemisphere fauna will undoubtedly result in new higher taxa being added and added to their proposed classification system.
Mitchell et al. (2006) reviewed the history of molecular sequence analysis of noctuoid relationships, the first of which was that of Weller et al. (1994) using partial sequencing of nuclear 28S rRNA (300 bp) and mitochondrial ND1 (320 bp) from twenty-six noctuoid species, including ten noctuids. They noted that despite low levels of support, parsimony analyses consistently grouped quadrifine noctuids with Arctiidae, and often Lymantriidae, rather than with trifine noctuids, suggesting paraphyly of Noctuidae and underlining the lack of unambiguous synapomorphies for this family. Friedlander et al. (1992, 1994), Mitchell et al. (1997, 2000), Fang et al. (2000) and Weller et al. (1994) all searched intensively for protein-coding nuclear sequences more suitable for resolving deeper phylogenetic relationships in Lepidoptera, mostly Noctuoidea (Mitchell et al. 2006). Two of the genes identified as being most useful were elongation factor-1α (EF-1α) and dopa decarboxylase (DDC), genes that have now provided strong evidence for relationships among some subfamilies and families of Noctuoidea. The study of Mitchell et al. (2000), which sampled a total of almost 2000 bp of DNA sequence from each of 77 species, provided strong evidence for noctuid paraphyly and for the monophyly of the true cutworms (Noctuinae s.l.).
Trifine and Quadrifine Noctuidae
Trifine refers to a large clade of noctuid subfamilies in which vein M2 in the hindwing is usually vestigial (Fig. Trifine noctuid with reduced M2 in Mamestra, Hadeninae or absent (Fig. Trifine noctuid with absent M2 in Peridroma, Noctuinae: Lafontaine & Fibiger 2006) so that the cubital vein appears to branch into three veins. In contrast, quadrifine refers to subfamilies in which vein M2 in the hindwing is as strong as vein M3, so the cubital vein appears to have four branches. In basal groups of quadrifine Noctuidae, vein M2 arises in the same position as it is in trifine Noctuidae, near the middle of the discal cell (Fig. plesiomorphic quadrifines hindwing in Chytolita, Herminiinae: Lafontaine & Fibiger 2006), whereas in more distal groups (apomorphic) the vein is adjacent to vein M3 (Fig. apomorphic quadrifines hindwing in Melipotes, Catocalinae: Lafontaine & Fibiger 2006), which according to Lafontaine and Fibiger (2006) is the true quadrifine hindwing venation. In addition, the families Nolidae, Strepsimanidae, Arctiidae, and Lymantriidae also have a true quadrifine hindwing (Fig. true quadrifine position in Spilosoma, Arctiinae: Lafontaine & Fibiger 2006).
Classifications of Noctuidae
In 1975, Nye (1975) stated that, “It is exceptional to find any two authors who use the same combination of subfamily names within Noctuidae”. Ten years later, Kitching (1984), in his historical review of noctuid subfamily relationships, commented that “The classification of the family is still in a state of uncertainty, and few authors appear to hold similar views regarding the suprageneric taxonomy”. His synopsis showed that the higher classification of Noctuidae at that time was based on superficial resemblance and vaguely defined characters, rather than on the rigorous application of cladistic principles. In an attempt to progress beyond the age of traditional noctuid taxonomy, Speidel et al. (1996) undertook an explicit cladistic study based on a small suite of morphological characters, such as the hood of the A1 pleuron, alula, membranous conjunctiva, tympanal bar, tympanum, male genitalia and their associated muscles, abdominal brush-organ, spur length and ventral cervical gland in the larva. Kitching & Rawlins (1998) used of character systems from both adults and immature stages for the first time in presenting the most thoroughly defended and comprehensive reclassification of the superfamily since Hampson´s (1898-1913) world catalogue (Fibiger & Lafontaine 2005). Although Poole´s (1995) classification was limited to the trifine Noctuidae (= Noctuidae s.s. of Fibiger & Lafontaine 2005), it has provided a basis for subsequent discussion of trifine noctuid classification (Lafontaine and Fibiger 2006). Beck (1999-2000) split Noctuidae into a large number of subfamilies based almost exclusively on larval characters, and recognized a very large number of family-group names, of which 68 were proposed by Beck either in this work or in a previous checklist (Beck 1996). These family-group names were diagnosed but not organized into a hierarchical classification on the basis of derived character states (Lafontaine & Fibiger 2006). The classification of Mitchell et al. (2006) is the most recent of three studies of Noctuoidea based on nuclear genes, but included twice as many taxa (146 species) as the two previous studies (i.e., Mitchell et al. 1997, 2000), and emphasized trifines, so that the authors gave a more definitive interpretation in terms of proposing changes to the classification than they had previously.
Fibiger & Lafontaine (2005) presented a comprehensive contribution towards a consensus classification of the Eurasian and North Americain faunas and synthesised their results with earlier morphological data and current molecular data to try to alleviate the current state of confusionof confusion. They generally followed Kitching & Rawlins´s proposed system but some fundamental changes were proposed (discussed in detail elsewhere). It is sufficient here to note the following changes that were related with the fate of the quadrifines:
- Placing the arctiid group of families (Nolidae, Arctiidae and Lymantriidae) in front of the upgraded family Erebidae so that their close relationship with the quadrifines is better reflected;
- Removing Lymantriidae from a position in front of Nolidae to a position after Arctiidae to reflect the close association of the arctiids and lymantriids following Mitchell et al. (1997, 2000);
- Treating the quadrifine subfamilies (Herminiinae, Hypenodinae, Hypeninae, Catocalinae, Calpinae, Hypeninae, Stictopterinae, and Euteliinae) as subfamilies of the re-established family Erebidae and as the sister group to the arctiid families, and reinstating subfamily Rivulinae following Fibiger and Hacker (1991), Speidel et al. (1996), and Beck (1999, 2000), subfamilies Boletobiinae and Phytometrinae following Beck (1999-2000), and subfamilies Scolecocampinae and Erebinae on the basis of their data; Hypenodinae was reinstated for Hypenodes Doubleday and its relatives, and Strepsimaninae was treated as family Strepsimanidae;
- Revising the classification of the subfamilies Catocalinae, Calpinae, and Erebinae by moving the tribes Armadini and Aediini from Catocalinae to Acontiinae, and Tytini from Catocalinae to subfamily Metoponiinae in the Noctuidae;
- Following Fibiger and Hacker (2002, 2004) and removing tribe Eublemmini from Eustrotiinae and treating it as a basal subfamily of Erebidae;
- Araeopteroninae Fibiger was proposed as a new subfamily of Erebidae.
One year later, Fibiger & Lafontaine (2006) once again reviewed the higher classification of the superfamily on the basis of recent morphological and molecular studies, and proposed a more inclusive definition of the family Noctuidae, one that added Nolinae, Strepsimaninae, Arctiinae, Lymantriinae, and Erebinae as subfamilies of a more inclusive Noctuidae.
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