Nolidae

The concept of the Nolidae has been expanded recently (Speidel et al., 1996; Holloway, 1998; Kitching & Rawlins, 1998) to embrace the Sarrothripinae, Chloephorinae and Camptolominae as well as the Nolinae sensu stricto as treated here. The first two have been treated as noctuid subfamilies during most of the last century (e.g. Hampson, 1912; Kitching, 1984). The third, though associated with the other two by Mell (1943), has often been associated with the Arctiidae (e.g. by Inoue et al., 1982); its status was reviewed by Holloway (1988) in Part 6 of this series. The Nolinae have also often been associated with the Arctiidae.

A detailed account of morphology within the group and a review of its classification may be found in Holloway (1998), together with description of the complex sound-producing (tymbal) structures found in males of many groups, usually at the base of the abdomen, but also in single instances in the hindwing (Cossedia Walker) and the valve, tegumen (Nycteola Hübner) and saccus of the genitalia (Cacyparis Walker). Sound production is also encountered at the pupal stage, the pupa stridulating a row of beads or carinae on the anterior margin of A10 against the interior of the cocoon, which may itself be ridged, a phenomenon discussed in detail originally by Hinton (1948).

The monophyly of the group, discussed in the next section, and its separation from the Noctuidae, increasingly seen as a paraphyletic assemblage (Kitching & Rawlins, 1998; Mitchell, 1998; Holloway et al., 2001), is also being confirmed in molecular studies, though on the basis of, as yet, a very small sample of taxa (Mitchell, 1998; Mitchell et al., 2000).

Definition of the family

Holloway (1998) noted seven characters, two of the pupal stage and five of the adult, that define the family.

The most reliable feature, but one that is not known for any taxa that have not been reared, is the structure of the cocoon: boat-shaped, with a vertical exit slit, and a two-walled method of construction. The pupa lacks a cremaster, but this is also a feature of the Euteliinae, currently in the Noctuidae. Some probably plesiomorphic groups have the abdominal beading or carinae of the pupa mentioned above.

In the course of dissection for this work, it became apparent that two of the features noted by Holloway (1998) that provide support for the aedeagus in the male genitalia are equally reliable and obviously more practical. The aedeagus is supported by projections from the valves anteriorly into the abdomen: dorsally there is a transtilla, extensions from the base of the valve costa that fuse apically into a U- or V-shape (Speidel et al., 1996); ventrally, similar processes extend from the base of each sacculus to fuse with a central shield-like structure, referred to hereafter as the saccular shield. It is not clear whether the central component of this second structure is homologous with the juxta. The features can be appreciated particularly easily when dissecting Risoba Moore species. The saccular shield can be seen clearly in Figs 2, 9, 26, 168, 176, 180, 200, 212, 263, 308, 376, 394, 452, 456, 478, 497, 511, 522 and 542.

Speidel et al. (1996) noted a ventral displacement of the ventral part of genital muscle m.4. Kitching & Rawlins (1998) considered that the elongate, bar-shaped retinaculum of the male forewing, though shared with a few other noctuoid groups such as the Arctiidae and Aganainae, was, when combined with presence of scaling of the lower part of the clypeofrons, also diagnostic for the Nolidae.

A final feature that needs further investigation is the observation by Holloway (1998) that the patagia and tegulae easily become detached from the thorax when this is compressed in the process of spreading specimens that have been relaxed from a dry condition. The morphological basis of this is not clear.

Other features that are common in the family, but not necessarily unique to it or found in all component groups include: the development of a pair of strong lobes or apodemes on the basal margin of the male eighth tergite and sternite, particularly the tergite (not in the Eariadinae); the presence of raised scales on the forewing (mainly in the Nolinae, Sarrothripinae, Bleninae and Collomeninae).

Several genera that have traditionally been associated with the groups now incorporated in the Nolidae lack the defining features and should probably be excluded (Holloway, 1998). These will be discussed in the final part of the systematic account.

Groupings within the family

A tentative classification into subfamilies and tribes is beginning to emerge (Holloway, 1998; Kitching & Rawlins, 1998; Holloway et al., 2001), but there is great disparity in the size of the divisions, some, such as the Nolinae and Chloephorinae, being very large, and others being monogeneric, such as the Bleninae, Risobinae (excluding Baileya Grote) and Eariadinae. There are also several genera and small generic groups that do not fall readily into these categories but nevertheless show typical nolid features.

The Nolinae (see also Holloway & Miller (1995)) are defined essentially on larval features such as reduction and loss of the first pair of prolegs, and presence of secondary setae on verrucae. The adults lack ocelli; these are present in the rest of the family and in Beana Walker, possibly the most plesiomorphic noline. There are also features of the male and female genitalia common to many genera.

The Chloephorinae assemble a number of tribes where at least some genera have tymbal organs at the base of the male abdomen. These show their greatest development and ubiquity in the Chloephorini (Figs 208, 211, 212, 214, 215) and Camptolomini, but also occur in all Careini (Figs 228, 230-239). In the Ariolicini (Figs 360-364, 396) and the Sarrothripini (Figs 126-131) they are present in only a few genera, though others have structures on the male basal sternite that may represent vestiges. These last two tribes are the most weakly defined. The Ariolicini have two subgroups, genera of the first mostly having peg-like setae on the valves or strong hair pencils at the valve bases, and those of the second having male genitalia bearing a possibly superficial resemblance to those of some Careini. The Sarrothripini frequently have a process on the valve that bears dark scales or setae, though this is not seen in the type genus, Nycteola Hübner. They usually have rather grey forewings with raised scales. Females of many genera have acute ovipositor lobes. These two tribes are the only ones in the Chloephorinae where pupae with beading have been noted.

No strong signals of relationships have been located in the head, wing venation or female genitalia (Holloway, 1998). The male antennae are generally fasciculate (filiform, with cilia), though bipectinate ones are seen in many Nolinae, two species of Labanda Walker in the Ariolicini and the Gelastocera Butler generic group. The absence of ocelli in Nolinae has been mentioned earlier. The labial palps generally have the third segment much shorter than the second, exceptions being discussed under Cacyparis Walker, which, with Ballatha Walker, has the third segment very long and slender. Reduction of the tongue is seen in several genera of the Ariolicini and some Collomeninae.

The groundplan venation is typical of the collection of groups currently assigned to the quadrifine Noctuidae subfamilies. The forewing has an areole, often elongate, at the anterior angle of the cell; this areole gives rise to the radial sector veins, usually with R2 and R5 arising independently from it on either side of the stalked (R3, R4). The hindwing typifies the quadrifine condition, with M2, M3, CuA1 and CuA2 arising independently, but often with M3 and CuA1 connate, around the posterior angle of the cell.

In a number of nolid groups this groundplan becomes modified. Stalking and reduction of hindwing veins is seen in the Nolinae, Sarrothripini, Ariolicini (loss only in Titulcia Walker, but an unusual branching system in Chandica Moore) and Eariadinae, and to a lesser extent (occasional stalking of M3 and CuA1)in the Chloephorini and Careini; in Phaeothripa Hampson and Ptisciana Walker of the Collomeninae M2 and M3 are stalked. Variations on the forewing groundplan consist of reduction and loss of the areole, and variations on the radial sector branching pattern, for example, with R5 branching from Rs more distally than R2 in some Sarrothripini and independently from the cell in others. In the Eariadinae and Chandica and Cossedia Walker in the Ariolicini R2 is independent but not R5. Radial sector veins are lost in some Nolinae such as Nola Leach.

The male genitalia show the aedeagus support structures that define the family. Reduction of the uncus is common, and there are often scaphial thickenings on the anal tube. The tegumen may be elongate, ventrally swollen on each side, extending beyond the junction with the vinculum and often bearing hair pencils (culcita of Kobes (1997)). The Chloephorinae and some other groups frequently have a subbasal process on the valve costa. A saccular harpe or other structure is less common, but is seen in the Nolinae, Risobinae, Bleninae, Westermanniinae and the genera Selepa Moore and Bryophilopsis Hampson. The valve is often paddle-shaped, with basally directed hair-setae distally on the paddle, which may be bilobed; these features are seen particularly in the Careini and the second group of the Ariolicini.

In the female genitalia the ovipositor lobes can vary from rounded, ring-like (many Ariolicini) to acute (many Sarrothripini). The sterigma is not often modified, and the ductus bursae is very variable in length. The occurrence and form of the signum or signa is also diverse, and general scobination of the bursa is also frequent.

In four widely separated genera (Mniothripa Hampson, Tympanistes Moore, Westermannia Hübner, Didiguides Kobes) the cornuti on the aedeagus vesica are deciduous, become detached during copulation and can be found in the corpus bursae of a mated female (Figs 192, 286, 299, 484). In all these cases the cornuti are somewhat dagger-like. This phenomenon is not unique to Nolidae but also occurs in, for example, ennomine Geometridae (Holloway, 1979: 338) and Notodontidae (Holloway, 1983: 4).

Early stages and host-plant specialisations

The larvae show a great range of features, including proleg loss and development of secondary setae on verrucae in the Nolinae as mentioned above. Much basic work on the chaetotaxy and other features was done by Gardner (1941, 1946a, b, 1947, 1948a) as reviewed by Holloway (1998) and Kitching & Rawlins (1998).

The bisetose/unisetose condition of the subventral setae on thoracic and bisetose/trisetose condition of this group on abdominal segments are variable within the family, but these features have been used to distinguish Noctuidae from Arctiidae (T unisetose, A bisetose versus T bisetose, A trisetose respectively). Thus Selepa and Earias show arctiid features, and the Camptolomini have an arctiid thorax but a noctuid abdomen. All other groups show the noctuid condition throughout. Other features are discussed under individual groups.

Pupal characters and the method of cocoon construction have already been discussed as defining the family; see also Gardner (1948b).

Some host-plant specialisation is seen in the family, or a non-exclusive focus on a particular plant family. Both the Nolinae (Nola Leach) and the Sarrothripini show a trend towards feeding on shoots, flowers, pods and seeds, and one noline at least has been noted as a gall feeder (Ito & Hattori, 1982). In the Careini, the Carea Walker complex of genera shows a high level of feeding on Myrtaceae, and Aiteta Walker is recorded only from the Combretaceae, particularly Terminalia. Terminalia and Combretaceae are also the only hosts noted for Westermannia Hübner and Miaromima Meyrick in the Westermanniinae. Several records for Blenina Walker and Labanda Walker are from Diospyros (Ebenaceae), but these genera have also been noted from other families such as Euphorbiaceae for the former and Dipterocarpaceae for the latter. Some Collomeninae, such as Gadirtha Walker, appear to be specialist on Euphorbiaceae. The Eariadinae show a temperate/tropical split between Salicaceae and Malvales. Some of the ‘careine’ sequence of the Ariolicini also appear to favour the Malvales.

Biogeography and ecology

The family is much more diverse in the Old World, particularly the Oriental tropics. Only the Nolinae, Sarrothripini of the Chloephorinae and Collomeninae extend to the New World, although Baileya Grote has been associated with the Risobini by Forbes (1954).

The matrices in Tables 1-3 provide biogeographic versus ecological profiles of the major groups recognised in the Nolidae. Whilst the Ariolicini and Careini show the high levels (around 40%) of endemics and Sundanian species with a preference for lowland forest that are characteristic of a high proportion of Bornean moth groups, the other two large groups, the Nolinae and Sarrothripini, include many more widespread species that tend to have a higher ecological amplitude, being found in both lowland and montane forests. The largest genera of the Nolinae are Nola Leach and Manoba Walker. Whilst the former shows the general character of the group, the latter has a higher number of endemic taxa, most of which are montane, though this endemic character may be an artifact due to the lack of data from other mountainous parts of Sundaland.

Amongst the smaller groups, the Collomeninae and the Westermanniinae show the more widespread type of profile, whereas the Risobinae, in the form of the genus Risoba Moore, have a greater proportion of endemics and lowland forest species, though many of these fall within one lineage of the genus.

There are three large genera in the Careini, Calymera Moore (21 species), Xenochroa Felder (36 species) and Didigua Walker (19 species), that all show the general Sundanian and endemic character of their tribe, though the first two contain more montane species and the last, with over half its species restricted to lowland forest, has a high proportion of those recorded from heath forest and forests on acid soils generally.

The sampling by Chey (1994) in softwood plantation and secondary forest habitats indicates that the group as a whole maintains a moderate to high diversity in these systems, particularly the Careini, Risobinae and Westermanniinae, and, to a lesser extent, the Ariolicini and Sarrothripini. The ability of the group to persist in cultivated areas is less clear, but probably not marked, except possibly in some Sarrothripini and Selepa Moore. This lack of persistence in more open and cultivated areas is probably a reflection of the almost entirely arboreal larval feeding by the group.

Strong specialisation in larval host plant preferences is not common. However, some groups appear to favour certain plant taxa, though rarely exclusively so. Feeding on Fagaceae is frequent in the Chloephorini and Camptolomini, particularly the more montane genera; montane Manoba species have also been recorded from Fagaceae. Many careine genera show a high number of records from the Myrtaceae, though Aiteta Walker species have been recorded almost entirely from Terminalia in the Combretaceae, a host also favoured by Westermannia Hübner and allies in the Westermanniinae. Many of the host records for the more advanced quadrifine Nolinae are also from the Combretaceae, and the family is exploited by Risoba, along with the Melastomataceae and several other families. Blenina Walker and Labanda Walker are often recorded from Diospyros in the Ebenaceae, and several genera in the Collomeninae appear to be associated with the Euphorbiaceae. The Malvales are favoured by Gariga Walker and allies in the Chloephorini, Paracrama Moore and Austrocarea Holloway in the Ariolicini, and by the tropical Eariadinae generally.