Edwards, in Nielsen et al. (1996), followed Kitching (1984) and Common (1990) in treating this group as a family and this was also supported in the molecular analysis of Mitchell et al. (2006). However, for reasons given on p. 4, the group will be treated at subfamily level here.
The group has been defined traditionally (e.g. Kitching, 1984) as consisting of quadrifine noctuids with a prespiracular counter-tympanal hood. This feature was recorded for a high proportion of taxa sampled from the group by Randolf & Lödl (2001). However, two definite herminiines were found to have a postspiracular, if rather distal, hood. These were from genera known from Borneo: Bocana Walker and Hadennia Moore. A third was Rhapsa Walker, a monobasic New Zealand genus referred to the Ophiderinae by Nye (1975) but placed in the Hypeninae by Dugdale (1988). Dugdale also noted that the counter-tympanal hood was postspiracular in both Rhapsa and Trigonistis Meyrick (see p. 222 for further discussion of the latter genus), and commented on the unsatisfactory nature of this character as a definitive apomorphy for Herminiinae, referring also to Bocana and other southern Pacific genera. This character is also labile in the Mecistoptera Hampson group of genera, treated under the Hypeninae on p. 160 et seq., and see also comments on the Trisatelini on p. 23.
However, Owada (1987) considered that a combination of this character of the tympanal organ, together with occurrence of a number of unique male secondary sexual organs and a feature of the male genital musculature, defined the Herminiinae as a natural group; see also Kitching & Rawlins (1998: 372), who also noted the feature of a swollen metepimeron associated with the tympanal organ. The male sexual specialisms are indeed diverse, but some are general to the whole group.
Perhaps the most consistent features are to be found in the female genitalia. The ductus bursae is robust but often long and reinforced by longitudinal strips of sclerotisation, most frequently two, these often rendering the ostium somewhat bilobed. Development of structures around the ostium is rare, but is seen particularly in Hydrillodes Guenée and Progonia Hampson; in Adrapsa Walker the ostium is often set in a pouch. The corpus bursae is generally elongate, sometimes more pyriform, and usually has extensive fields of scobination centrally and / or distally. In some genera this may be concentrated into signa, and there is also a generic grouping where the scobination is coarse and is distributed in a horseshoe shape in the distal half of the bursa. The basal section is often sclerotised and may lack scobination; it may also be corrugated. The ductus seminalis arises from somewhere between the base and the centre of the corpus bursae, most frequently subbasally. It often forms a small appendix bursae, tapering away from the bursa, and is usually coiled to a varying degree from a partial turn to several. The latter extreme is illustrated particularly by Hydrillodes where there are many rather tight turns.
Owada (1987) made a particular study of the secondary sexual characters of the males, these occurring as modification of the antennae, labial palps, forelegs and wings. The antennae are usually ciliate or fasciculate, there often being a pair of larger and more robust bristles to each segment. More rarely are the antennae bipectinate. In several genera, but not necessarily all species thereof, the antennae are noded between one third and half way along the flagellum. The node is formed in various ways, as illustrated by Owada (1987), and may involve a combination of shortening with expansion of the flagellomeres, twisting, and development of very robust hooked bristles from setae on two or three flagellomeres. Occurrence of nodes is scattered through the subfamily, such as in Paracolax Hübner and Hadennia Moore in one lineage, and in Nodaria Guenée, Simplicia Guenée and Polypogon Schrank in another, and also Adrapsa outside these lineages.
The labial palps are generally prominent, recurved, rarely straight. In a number of genera, many probably related to each other, they become massively extended over the head to reach the thorax or the anterior part of the abdomen. They may be held at rest in grooves along the dorsum of the thorax in extreme cases. Often there are hair-pencils associated with the junction of the second and third segments. Extreme examples may be found in Catadoides Bethune-Baker, Mosopia Walker, Oxaenanus Swinhoe and Echanella Bethune-Baker. In some genera where they do not extend beyond the head, they can nevertheless be densely tufted with scales in the male (e.g. in Hydrillodes).
The forelegs in many genera have unique tibial sheaths, the tibia becoming shortened from its junctions with the femur and first tarsal segment, but at the same time extending in the form of a sheath to cover much of the latter. In one lineage the proportions of the various segments can vary dramatically, with the coxa, trochanter, femur and first tarsal segment becoming relatively long, some extremely so. Some tarsal segments may become very small or lost and, in several genera, the tarsal system is reduced to a spine-like first tarsal segment. Tufts of hair scales arise from the femur, the tibial sheath and the first tarsal segment, presumably used in courtship, leading to the English vernacular name for the group as ‘fanfoots’.
Further variations on this theme involve the development of processes at the junction of the first and second tarsal segments, usually at the distal end of the former. Sheaths are found in two major generic lineages and also in genera found further east in Fiji (Robinson, 1975).
Forewing modifications are seen in males of several genera, the commonest being a scaled subcostal flap. It is illustrated for some Japanese species by Owada in Inoue et al. (1982: plates 383, 384). This is narrow and not very conspicuous in Hydrillodes, Echanella, Adrapsa and Bocana, but is welldeveloped in Paracolax and Hadennia, probable sister-genera, and in Raphiscopa Hampson and Ochrotrigona Hampson. Unique modification to the upperside of the wing, such as patches of androconial scales, flaps or pockets are seen in Mosopia, Globosusa Swinhoe, Echanella and Hydrillodes. In Hydrillodes, some groups have a large semicircular flap at the centre of the forewing costa that covers a hair-pencil or a fringe of hairs.
The wing pattern is diverse across the group, but there are two features of the hindwing that are unusual and may serve to identify subgroups within the family. The first is an obtusely angled submarginal, noted by Holloway, Bradley & Carter (1987) and Holloway et al. (2001), and this occurs most frequently in an extreme form in Simplicia and related genera where the corpus bursae of the female contains a horseshoe arrangement of scobination, as discussed above. The second feature is the presence on the underside of a small dark spot basal to a similarly dark discal mark, effectively an orbicular stigma. This is widely distributed across several of the genera with strongly developed labial palps, but also occurs in a species tentatively described in Subsimplicia Prout (p. 107). The forewing usually lacks an orbicular stigma.
The male abdomen has the eighth segment of the framed corematous type in many genera. Where the corema of the sternite is developed, it is usually single, central, a feature shared with the Aganainae. If loss of the framed corematous state is treated as apomorphic, as for the Catocalinae (Holloway, 2005: 14-15), then this is another feature shared by the genera with a hindwing orbicular and well developed male labial palps and is present also in a number of other genera outside the Simplicia group.
In the genitalia, the uncus tends to be well developed, deep, sometimes very much so over some part of its length, but tapers to a small, usually downturned point apically; some species have the dorsal part domed or with a spine. At least the dorsal part is generally setose. The tegumen is usually simple, rarely with processes (but see Hydrillodes on p. 75), and the vinculum is often broad but may extend and narrow into a saccus. The juxta is platelike, often triangular, with the apex dorsally. The valves are often simple, tongue-like, modification being mostly in the form of distal extension of the costa or sacculus into various processes or spines, and sometimes such processes occur more basally on the margin. Again Hydrillodes is unusual in having the valves distinctly bifid. The apical part of the valve may also become narrowed or acute, sometimes leading to a trifid arrangement as in many Japanese Zanclognatha Lederer (Owada, 1987, and see p. 124). In Paracolax and Hadennia there is a spherical structure between the bases of the valves that gives rise to a hair pencil (see p. 87). The aedeagus is usually a straight or slightly curved tube that may have scobination or a few spines apically. The vesica is large, characteristically angled dorsally (or in the same direction as the insertion of the ductus ejaculatorius), from the axis of the aedeagus. It is usually strongly lobed but rarely has diverticula that are very long. Scobination tends to extend over most of the vesica, but the spines may vary considerably in size, there often being small fields of large, blunt ones, and sometimes these may fuse into a spiny sclerite. Large cornuti are rare and tend to be basal when they occur. These vesica characteristics are general to the subfamily and may provide a further diagnostic feature.
In general, the species are of delicate build. The legs are relatively longer than in most other noctuoid groups. The phragma lobes between the first and second abdominal tergites are large, tongue-like.
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