SUBFAMILY NOLINAE
View Image Gallery of Subfamily Nolinae

The classification of the Nolinae has been discussed by Holloway & Miller (1995), Holloway (1998) and Holloway et al. (2001).

Most of the species are small, the forewings variegated black, grey and white, often with areas of raised scales; more colourful species are seen in a few genera such as Evonima Walker. The male antennae are filiform and ciliate or bipectinate, the former probably being the plesiomorphic state, and possibly secondarily derived in Evonima. Those of the female are filiform except in Dialithoptera Hampson where they are strongly bipectinate in both sexes. Ocelli are weak or absent, strongest in
Beana Walker, possibly the sister-genus to the rest of the subfamily.

The forewing venation usually lacks an areole except in Barasa Walker and allies, though a small one may occur in more advanced taxa such as some Meganola Dyar. The radial sector veins posterior to R1 may be reduced to three or two from four, and this was used by Hampson (1900) as the basis for his classification. Reduction from three to two occurs within the more strict definition of Nola Leach of Holloway & Miller (1995). The hindwing is of the Nolidae groundplan type in more basal genera, but with M3 and CuA1 stalked in all except
Beana, and M2 separated from this. A trifine condition arises in more advanced genera (mostly containing smaller species), probably through fusion of the stalked veins as there is still a separate vein in the M2 position.

The male abdomen lacks tymbal organs but usually has apodemes on the eighth tergite, and these may also occur on the sternite in more derived genera where both sclerites can become distinctly narrowed. The genitalia have a simple uncus and simple valves with a characteristically ‘noctuid’ saccular harpe. The saccular shield is often small, modified or lost, particularly where the saccular bases meet broadly in the centre of the diaphragma. The most characteristic feature seen in many genera is a pair of setose lobes that flank the scaphial area of the anal tube (e.g. Figs 2, 7, 70). Extreme modification of the uncus is seen in Beana and
Ctenane Swinhoe where it gives rise to rake-like structures, though these are located differently and possibly not homologous. The uncus is lost in most Nola, which also have the valve diagnostically cleft. The aedeagus can have apical projections, and the vesica may contain cornuti.

 




The female genitalia have no particularly diagnostic features in segments 8-10, except the ovipositor lobes are usually short, quadrate or, seen together, rather conical. The ductus and neck of the bursa are often slender, elongate. There may be a small appendix bursae. The corpus bursae itself is usually pyriform and contains signa, usually two, opposed scobinate corrugations (Fig 6) or bands in more primitive genera, and invaginated ridges or thorn-like structures in more advanced ones (e.g. Figs 30, 86). Dialithoptera is unusual in having two stellate signa similar to those of ennomine geometrids (Fig 18).

 



The larva is characterised by a reduced anterior pair of abdominal prolegs and, in all known larvae except that of Beana, secondary setae on verrucae. An unusual feature recorded for a number of genera and species is the retention of a stack of old larval head capsules in a horn on the ‘current’ head (McFarland, 1980; Holloway & Miller, 1995; Holloway, 1998). This feature is recorded amongst more derived members of the quadrifine sequence but is not known from trifine genera except Evonima and one example in Manoba.

The generic classification within the subfamily is far from satisfactory (Holloway & Miller, 1995), with both nomenclatural confusion and a lack of a clear understanding of morphological features. Genus-group names such as Nola, Celama Walker and Roeselia Hübner have been applied indiscriminately in the past (e.g. Hampson, 1900; van Eecke, 1926), with other genus-group names assigned as synonyms in a similar fashion; Celama and Roeselia in fact fall within a strict definition of Nola, leaving a considerable number of both quadrifine and trifine genera outside this definition (old, erroneous concepts of Roeselia and Celama respectively). The quadrifines were placed in Meganola Dyar and the trifines retained in Nola by Poole (1989), and both are currently becoming assigned to the older name Rhynchopalpus Hampson by workers such as Inoue (1998, 2001) following Holloway & Miller (1995). The trifine taxa are reconsidered under Manoba Walker, and other trifine genera are discussed, particularly in the Australian and African faunas.

 

Fig 1. Tentative phylogeny for the Nolinae as discussed, with indication of possible apomorphies as follows: A, loss/reduction of first pair of larval prolegs; pair of scaphial lobes present in male genitalia (also elsewhere in Nolidae); B, larva with verrucae; C, male antennae bipectinate; D, female with invaginate signum or signa in bursa; larval head-capsule stacking widespread; E, hindwing venation trifine; F, partial or complte loss of larval head-capsule stacking.

The classification emerging for the group from current studies, including this one, is a rather pectinate one as summarised tentatively in Fig 1. Beana is basal for reasons given earlier, followed by the Barasa group and Melanographia Hampson, quadrifines with M3 and CuA1 stalked, primitive corrugate signa and filiform male antennae. The next sequence of genera is marked by possession of bipectinate male antennae and development of the invaginated type of signum, the latter not seen in Dialithoptera, which is therefore placed in the most basal position, below an unresolved complex of quadrifine genera with modified signa and male antennae bipectinate. It is amongst these that most records of larval head capsule stacking occur: the Acatapaustus Bethune-Baker (= Eurynola Hampson) complex in the Australasian tropics (Bigger, 1988); Uraba Walker (= Coesa Walker and Toxoloma Felder) in Australia (McFarland, 1980); Proneca Swinhoe and Sarbena Walker (see below); unplaced quadrifine species from the old concept of Roeselia/Meganola such as gigas Butler, mikabo Inoue, nitida Hampson and togatulalis Hübner (McFarland, 1980; Sugi, 1987).

Head capsule stacking does not provide a means of defining a natural group of quadrifine genera or a single genus, as records of it are too sparse, and it also occurs in Evonima where hindwing venation is reduced to the trifine condition and male antennae are filiform, possibly secondarily so, and in one species of
Manoba where the stacking habit is possibly more vestigial and the capsules less tightly associated. Thus these quadrifine taxa form an unresolved polychotomy in Fig 1, and generic placement of such taxa is problematic as there are no obvious features that permit assignation of quadrifine "Roeselia” to the genus-group names already available or justify description of new genera, hence these species are placed in "Meganola”. Features that may be helpful in resolving this sequence of quadrifine taxa and its relationship with the trifine sequence discussed next include: a longitudinal arrangement of the two signa in the bursa, seen in several "Meganola” species  (Figs 30, 32, 38, 41; see also illustrations for some Meganola species in Inoue (1982)), but also in Evonima (Fig 37) and brunellus Hampson and allies in Manoba; a double-peaked area that stains darker with chlorazol black on the basal margin of the male eighth sternite, seen in Figs 29 and 36.

 


Evonima
, with head-capsule stacking, is placed as sister-group to the remaining trifine taxa where weak or no head-capsule stacking has been recorded and where it has probably been secondarily lost. The relationships of remaining trifine genera are currently unresolved. Nola can be clearly defined, and a concept of Manoba Walker, an older name, is offered below for the bulk of trifine
Rhynchopalpus / Meganola taxa. However, there are additional trifine genera in Australia (Dimona Walker, Sorocostia Rosenstock, Tribunta Walker), Africa (e.g. Metanola van Son, Paranola van Son, Poliothripa Hampson) and the New World (e.g. Callinola Butler) where the relationships are unclear except they do not fall within the definitions of Nola and Manoba given here.

Limited material of several other Bornean noline species was also examined in London and from the Zoological Museum, Univ. Amsterdam. These species could not be assigned with confidence to any of the generic concepts discussed below. Discussion of them is therefore held over until more material becomes available and the studies of colleagues in the Hungarian Natural History Museum, Budapest, and their associates have established clearer generic concepts based on an appreciation of taxa from throughout the range of the subfamily. Reference to this research effort and information kindly provided to the author will be attributed to the 'Budapest group'.

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