L.A.Q. CLADE
Weller et al. (1994) showed, surprisingly, that the quadrifine examples, including a core catocaline (Catocala Schrank), a core calpine (Plusiodonta Guenée) and a hypenine (Plathypena Grote), all grouped with Arctiidae and Lymantriidae rather than with the group of trifine examples in analyses using combined DNA sequence data. These results raised serious questions about the status of Noctuidae by indicating that the family was not monophyletic, implying that the cutworms are a grade rather than a clade within the superfamily. Weller et al. (1994) used the mitochondrial gene ND1 and a nuclear ribosomal RNA (rRNA) data set in their phylogeny estimations. They analyzed the mtDNA data separately under different assumptions of character treatment, such as unweighted sites, transversion parsimony, and third codon site omitted, before combining the two data sets.
The gist of Weller´s discovery was that the trees based on the combined data matrix were in closer agreement with previously proposed morphological trees (Fig. Traditional Phylogeny Tree of Noctuoidea), although there were striking differences regarding the placement of the quadrifine noctuids (Fig. Strict consensus tree resulting from unweighted analysis of combined rRNA & mtDNA sequence data & Fig. Tree resulting from SACW: Weller et al. (1994)). As in the mtDNA trees, the quadrifine noctuids remain embedded within Arctiidae, and Lymantriidae is the sister group to Arctiidae/quadrifine noctuids in only half of the trees.
The mol ecular analyses of Mitchell (1998), Fang et al. (2000) and Mitchell et al. (2000) only included three taxa from the Catocalinae / Ophiderinae complex. Two were core catocalines, a species of Catocala Schrank and a Caenurgina McDunnough from the Euclidiini. They selected the type species of Hypsoropha Hübner as the representative of the Calpinae, a genus that was also included in his concept of this group by Fibiger (2003), although Holloway (2005), who had circumscribed the taxon, noted a number of unusual features that made its placement unclear.
Holloway (2008) noted that several molecular studies had recently indicated that the traditional concept of family Noctuidae was not monophyletic. If monophyly was the most important criterion, then the family should be restricted to the trifine groups covered by Holloway (1988), with the further inclusion of Plusiinae, Pantheinae and most of the traditional Acontiinae / Eustrotiinae complex, as suggested by many other authors (i.e. Weller et al. 1994; Mitchell et al. 2000 & 2006; Fibiger & Lafontaine 2006). However, this left out virtually all the quadrifine subfamilies of the traditional concept of Noctuidae. Molecular results showed these intermingled with Lymantriidae and Arctiidae, with some surprising associations, such as a sister-relationship between Aganainae and the noctuid subfamily Herminiinae (Holloway 2008). Mitchell et al. (2006) illustrated for the first time some detail of L.A.Q. monophyletic relationships, which placed nearly all quadrifine noctuids (Erebidae sensu Fibiger & Lafontaine 2005) plus Arctiidae and Lymantriidae in a clade, which they termed the L.A.Q. (Lymantriidae, Arctiidae + quadrifine Noctuidae) lineage. There was very strong support (BP >= 90%) for this L.A.Q. Clade (Fig. Strict consensus for the 146 taxon: Mitchell et al. (2006)).
Lafontaine & Fibiger (2006) defined the L.A.Q. lineage as a clade containing Lymantriidae, Arctiidae and part of the quadrifine Noctuidae (i.e. Hypeninae, Aganainae, Herminiinae, Calpinae, Catocalinae, Scolecocampinae [as Strepsimaninae (Phobolosia Dyar) in Mitchell et al. 2006], Boletobiinae [as Rivulinae (Mycterophora Hulst) in Mitchell et al. 2006], Eublemminae [as Eublemmini in Mitchell et al. 2006], and Rivulinae (s.s.)) (Fig. Rivised classification of Noctuoidea presented by Lafontaine & Fibiger 2006).
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