::The Moths of Borneo::

SUBFAMILY BAGISARINAE

This subfamily was erected by Crumb (1956) primarily on the basis of an unusual larval feature, the presence of appendiculate (toothed) crochets on the prolegs. Crumb suggested that the Old World genus Xanthodes Guen�e might also be associated. Holloway (1998), Kitching & Rawlins (1998) and Sugi & Sasaki (2001) explored the limits of this concept further and concluded that most of the taxa attributed by Gardner (1946a, 1947, 1948a) to his group B1 on the grounds of possession of appendiculate crochets should be included in the Bagisarinae.

�� The included taxa all have prolegs on A3 and A4 vestigial, reduced to small tubercles or cylinders; those excluded are in the genera Cosmophila Boisduval and Marcillada Walker, where the prolegs on A4 are well developed, both treated by Holloway (2005). The larvae are thus highly active semi‑loopers with a leech-like motion, particularly in early instars, when they tend to live on the undersides of leaves. An escape reaction recorded in many of the genera is for the larvae to wriggle and jump off, suspended by threads.

�� There is a strong, but not exclusive, association with host plants in the Malvales. Gardner (1948b) also drew attention to pupal similarities within the group in the form of a pair of straight cremastral spines arising from a rounded extremity, supported for further Indian species by Bell (MS), though the spines may be curved. However, these features are not unique to the group, as discussed later.

�� However, no really incontrovertible synapomorphies have been located amongst adult characters. Sugi & Sasaki noted a development of the male eighth tergite exhibited by many genera that is a modification of tergal structure that frequently accompanies the framed corematous condition of the eighth sternite. The apodemes of the anterior margins are closely associated and give rise to a central thickening or rod that extends into an abrupt disc-like or ring-like expansion that renders the tergite as a whole somewhat paddle-shaped or shaped like a table-tennis bat. Holloway (1989) described the feature in Dyrzela Walker as resembling the ring-pull of a drink can. The valves of the male genitalia, when spread, tend to have the costal margin adjacent to the tegumen, or not widely splayed from it, and the ornamentation of the interior side may be complex. The valve bases tend to be projecting as a lobe often with a hair pencil exteriorly, and may show some fusion together interiorly across the diaphragma. The tegumen and vinculum are often joined on each side by a diversely complex structure that suggests involvement of a paratergal sclerite or pleurite.

�� In typical Bagisara Walker, the male abdomen has the apodemes of the basal sternite with a slightly falcate flange interiorly. The eighth segment has a paddle‑like tergite, but the sternite is much enlarged into a sac‑like pouch with a single corema; its posterior margin extends into lateral wings that bear brushes of scales. The genitalia have the two valve bases fused with each other centrally. The valves are short, with spurs at the centre of the costa and at the distal end of the interior margin of the sacculus. The saccus is large, with an apical nipple. In both sexes, the phragma lobes between the first and second abdominal tergites are present and moderate in Bagisara, but much shallower in other genera.

�� The female abdomen offers no obviously autapomorphic features, but the genera Bagisara, Calymniops Hampson, Chasmina Walker, Elydna Walker (see below) and Xanthodes all have a mostly globular appendix bursae that arises on a narrow tube from the base of the corpus bursae; it may contain spines. The other genera included here have the ductus seminalis arising directly from the base of the corpus bursae or (Imosca Sugi & Sasaki, Ramadasa Moore) from a small, tapering appendix bursae in the same position. The ostium is situated close to the anterior margin of the eighth segment and sometimes fused with it. The corpus bursae is variable, sometimes small and flimsy, but, when large, can be extensively corrugated and / or scobinate. There is no signum, however.

�� The facies has some features that recur across several genera such as fine, often linear forewing fasciae with a well defined dark patch on the costa just distad to the postmedial. The scaling of the wings is often very fine, regular, even satiny.

�� The treatment of the subfamily here differs from that of Fibiger & Hacker (2007) and Hacker & Fibiger (2007) by including the genera Imosca and Brevipecten Hampson (and, by association, Honeyia Hacker & Fibiger) in the Bagisarinae rather than in the subtribe Cosmiina of the tribe Xylenini. This placement was apparently made �as indicated by Fibiger & Lafontaine (2005)�. However, the only mention of Brevipecten by Fibiger & Lafontaine (2005) is within the context of their Erebinae (p. 29), not the Cosmiini (p. 44). Here they stated that cosmiine larvae feed on tree foliage, tying several leaves together to form a feeding shelter. Fibiger & Hacker (2007: 148) stated that Lafontaine & Fibiger (2006) placed the Cosmiina within the Xyleninae �on account of the structure of the male and female genitalia, and especially on the structure of the larvae and their habit of feeding on foliage of trees�. Cosmiina larvae are typical of trifine noctuids, rather stout, with all prolegs fully developed. Therefore, on larval features and behaviour, Imosca and Brevipecten are clearly Bagisarinae.

�� The male genitalia structure of the Cosmiina, illustrated by Fibiger & Hacker (2007), particularly of the valves and the presence of a peniculus on the tegumen,� does not appear to be similar to that of Bagisarinae, though homologies of the somewhat complex paratergal sclerites in both groups need assessment. Fibiger & Lafontaine (2005) did not redefine the Bagisarinae except to note, in transferring the Cydosiini to the group, that the bases of the valves are broadly fused together; this feature appears to occur in Brevipecten as figured by Hacker & Fibiger (2007), and the valve shape and complexity is more as in some Bagisarinae; the valve bases in some species are externally expanded and sometimes corematous. The eighth segment of typical Cosmia Ochsenheimer males is unmodified. Some genera of Cosmiina have a spined apex to the ovipositor lobes (Fibiger & Hacker, 2007), but this feature does not occur in any genera attributed here to the Bagisarinae. Mode of pupation (in a cocoon in the soil) and pupal abdominal structure appears to be similar in Bagisarinae (see above) and Cosmiina (Bretherton et al., 1983), but the features mentioned are widespread in the trifine Noctuidae.

�� Four further Bornean genera are included in the Bagisarinae here. Calymniops resembles Dyrzela externally but differs somewhat in abdominal features, for example the large, coiled appendix bursae of the female; the early stages are unknown. Androlymnia Hampson was included in the original list of genera noted by Gardner (1946a). Ramadasa shows some adult features that indicate it may be bagisarine as discussed on p. 35 and suggested by Holloway (2005: 26). The larvae and pupae of species in Amyna Guen�e are of the bagisarine type (Gardner, 1946a, 1948b), and the genus is undoubtedly misplaced in the Acontiinae and Eustrotiinae as defined here. Some features of the male abdomen are also consistent with placement in the Bagisarinae as discussed on p. 38.

�� Brevipecten is not represented in Borneo, but six species occur in the Oriental Region, including the type species, captata Butler, and one in Australia; the centre of diversity for the genus is Africa where 23 species are known (Hacker & Fibiger, 2007). Sugi & Sasaki (2001) observed that the larva of B. consanguis Leech has been recorded as feeding on Hibiscus in China.

�� Gardner (1946a, 1948a) and Bell (MS) noted a number of other taxa in the Indian Subregion that have larvae with bagisarine features, though Bell did not note the condition of the crochets. Several of these extend into S.E. Asia and further east, or have more easterly relatives, but none is currently known from Borneo. Several are listed under Oglasa Walker by Poole (1989), as discussed by Hacker & Fibiger (2007), and three are listed by Poole under Athetis H�bner. One of the former, separata Walker, is tentatively combined with Ramadasa Moore on p. 36, comb. n., because of shared features of the male genitalia.

�� The taxa included in Athetis are here extracted from that genus and placed in Elydna Walker Gen. rev., one being the type species thereof, transversa Walker comb. rev. Holloway (1989, 1998) and Fibiger & Hacker (2007) indicated that the subordination of Elydna to Athetis required further study, and Gardner (1946a) stated clearly that the larva of transversa had the characteristics of his group B1 and was not related to the species of Athetis that he had also examined. Bell noted behaviour on disturbance typical of Bagisarinae. The host plant recorded by both Bell and Gardner was Schleichera (Sapindaceae), a member of the Sapindales rather than Malvales (Mabberley, 1987).

�� The male abdomen of E. transversa is unusual in that the apodemes of the basal sternite are flanked by lacunae or pockets that are lined by dense rows of scales on ridges, and the structures on the eighth segment are reduced to rod‑like apodemes. In the genitalia, the valves are fused at the base and show marked bilateral asymmetry, being divided into a paddle‑like dorsal, cucullar portion and a highly sclerotised ventral, saccular portion with lateral spurs and an internal hook.

�� The two other species listed by Poole in Athetis, but here placed tentatively in Elydna, are E. atripuncta Hampson comb. rev. (Sri Lanka, India) and E. ochracea Hampson comb. n. (Burma, Java, Bali, Philippines). These have ochraceous fawn or grey forewings with faint and highly irregular fasciation, and are sister‑species, having very similar male genitalia. Bell (MS) reared atripuncta and noted proleg reduction and escape behaviour in the larvae that is typically bagisarine. The host plant was Grewia (Tiliaceae) in the Malvales. In the male abdomen there is one possible synapomorphy with E. transversa in the development of a secondary sexual feature associated with the apodemes of the basal sternite, though in this case it is in the form of a pad‑like structure on an inturned lobe on each side exterior to the apodeme. The eighth segment is typically bagisarine. In the genitalia, the valves are partially fused at the base, expanding distad, and appear bifid distally through the occurrence of a prominent central process to the costal margin. There is a thorn‑like process distally on the interior of the saccular margin that may be homologous with the internal hook noted in E. transversa. This is less evident in ochracea.

�� Gardner (1946a) also identified �Oglasa� hypenoides Moore (Indian Subregion, Bali, Sumbawa) as having a larva with bagisarine features. It also feeds on Grewia. The facies resembles that of Imosca and typical Elydna, but lacks the dark patches of the former and has an additional straight medial fascia; the submarginal is crenulate. The male abdomen has a small anteriorly directed lobe exterior to the posterior (basal) end of each apodeme on the basal sternite. There is a pair of large, centrally conjoined pouches full of deciduous hair‑scales across the third tergite. The eighth segment is typically bagisarine. The genitalia have the uncus short, its apex of the ball-and-claw type, but only just protruding beyond an expanded, dorsally flat disc that bears crests of hairs. The tegumen is complex, sinuous, with elongate, hair‑bearing pads. The valves are basally fused, narrow, of bagisarine orientation, and divided into a shorter, very slender dorsal process and, extending from a broad sacculus, a narrow, longer, central process that extends to the level of the uncus, bearing on the interior of the apex, two appressed, slender spines. This species is hereby also transferred to Elydna, comb. n., on the grounds of general facies and the presence of structures on the basal abdominal sternite of the male, though this is probably only a temporary placement, but here serving to bring the species into the Bagisarinae.

�� Apocalymnia tenebrosa Hampson is an Indian species that typifies the monobasic genus Apocalymnia Hampson. It is a robust species with a finely irregular and variegated forewing facies. It has not been dissected, but Bell (MS) described proleg reduction, larval behaviour and pupal structure that is consistent with placement in the Bagisarinae. The host plant he recorded was also Grewia.

�� Hacker & Fibiger (2007) noted an African species, renilinea Gaede, that was listed in Oglasa by Poole (1989) but appeared to be more related to Brevipecten. It is distinguished in particular by a pair of long, slender, hook‑like socii flanking the uncus in the male genitalia which also have prominent �shoulders� to the tegumen. Hacker & Fibiger recognised similar species from Madagascar and Thailand, but did not identify either. The latter proves to be �Oglasa� pellicea Swinhoe, known from Thailand, Java, Bali, the Philippines and Sulawesi. A further species in this group, costiplaga Bethune‑Baker, occurs in New Guinea and is very similar in facies to pellicea. No information on early stages has been located, but this group is probably also bagisarine, having a typical structure to the male eighth abdominal segment.

�� With the species in the genera Dyrzela and Chasmina Walker covered by Holloway (1989), the Bornean fauna of the subfamily comes to 26 species. The biogeographic and ecological profile for these is presented in Table 1. Most are lowland species, though some of these extend up into montane zones. A few very widespread species are found in open habitats, particularly in the genus Amyna Guen�e, and these can be minor pests.

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