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Nola ?quadrimaculata Heylaerts stat. rev.
Nola quadrimaculata Heylaerts, 1892, Annls. Soc. ent. Belg., 36: 43.


Nola quadrimaculata

Diagnosis and taxonomic note.
This species belongs to a complex that has previously been attributed to internella (see above) but distinguished by Wileman & West (1928) in their description of
analis (Sri Lanka, India) and distinction of it from pascua Swinhoe (India); both pascua and quadrimaculata were placed as synonyms of internella by Hampson (1900). All three species show sexual dimorphism, the males with yellower hindwings that have hair pencils and patches of brown scales associated with the dorsum. In analis and quadrimaculata there is a hair-pencil enfolded in the dorsum that is lacking in pascua (Wileman & West, 1928) and all have brown scales on the underside at the tornus. As these scales can become worn and lost, it is advisable to confirm indentification from the genitalia. The forewing facies has the double postmedial with a subdorsal kink (particularly strong in pascua) referred to above with reference to internella. The submarginal is irregularly triarcuate, defined by a sharp boundary between a distal darkening and a basal lightening. There are conspicuous patches of dark brown raised scales on the costa medially and antemedially. The male genitalia differ particularly in the shape of the harpe and in the ornamentation of the aedeagus vesica as illustrated (analis: Figs 94, 95; pascua: Fig 92). The female genitalia are also diagnostic, with a pair of lobes or ridges of the lamella postvaginalis short, adjacent in analis (Fig 96), longer, basally separate in pascua (Fig 97), and widely separated, incurving grooves in quadrimaculata. The ductus is much longer in pascua than the other species, and the bursa is round in analis, an elongate pyriform in quadrimaculata, intermediate in pascua. In pascua there is a more basal invaginate signum and a more distal one of two scobinate patches. Only the latter occurs in the other species and is very much reduced in quadrimaculata. The taxon internelloides van Eecke (Sumatra) is a distinct species with males resembling females of the above complex. The type of quadrimaculata was not located by the author in RMNH, Leiden, and is stated in the original description to be a male from Java with greyish white hindwings, hence its status is unclear, and attachment of the name to the widespread species of the analis complex is tentative.


Geographical range.
Indo-Australian tropics from N.E. Himalaya to Queensland and the Solomons; the distribution in mainland Asia requires further investigation.

Habitat preference. The few records in recent surveys are from lowland localities with secondary forest, a singleton from an arboretum, two from coastal forest in Brunei and two from lowland alluvial forest near G. Mulu.

Biology. Given the confusion over the identities of species in this complex, it is difficult to disentangle all the biological data that have in the past been attributed to
internella or analis (e.g. in Robinson et al., 2001). The description of the biology and illustration of the larva and cocoon of internella (see above) by Hampson (1900) may refer to pascua or quadrimaculata. The head is brown, T1 and T2 are reddish and the rest of the body is yellow to A6. There are two broad black subdorsal stripes on A1 and A2, continued more narrowly back to A6, and a purple lateral stripe from T2 to A9. The anal area is reddish brown, and the verrucae bear short pale hairs. The cocoon is compact, boat-shaped.

The host plant was Rubus (Rosaceae), the larvae feeding inside the fleshy young shoots.

Bell (MS) reared what he considered to be analis in southern India. The larva is typically noline, the body light yellow with a lateral purplish band and ventral suffusion of the same colour. The lateral verrucae are yellowish, the more dorsal ones orange, white and black. A8 has a trapezium-shaped purple dorsal patch that attenuates anteriorly to a line forward to A4, but A1 to A3 are more broadly purple dorsally, and the thoracic segments also have purple marks. Most secondary setae are light grey or translucent, but some are orange on T1, T2, A8 and A9; the large dorsolateral verrucae on A1-3 and A7 bear black setae. The setae are variable in length; the longest can attain half the length of the body. The head is a glossy pale orange with darker marking. Bell also noted variants that were yellow and black with orange verrucae and a green ventral surface, and others brownish black, marbled with yellowish white, with purple, orange and white verrucae.

The eggs are laid singly or in batches. The larvae are restless, the early instars very hairy and sometimes blown away by wind. They live and feed on the flowers of the host plant. Pupation is in a triangular cocoon, peaked anteriorly, that incorporates bark particles and is attached to a twig or branch. The pupa lacks a cremaster, and is bluntly rounded at both ends.

Bell also described a larva that he considered might be pascua, with a glossy black head and a pinkish brown body, lined with white and yellow. The verrucae are yellowish except for the dorsolaterals of A1-3 and A7 which are shiny black, set in velvety black patches and bearing black setae (these are white on other verrucae); there is a dorsal black patch on each of T1 and A8.

The host plants recorded by Bell in southern India were Memecylon (Melastomataceae) and Terminalia (Combretaceae).

Other host-plants noted for the complex, mainly from India, are (Robinson et al., 2001): Mangifera (Anacardiaceae); Durio (Bombacaceae); Ricinus (Euphorbiaceae); Pennisetum, Sorghum (Gramineae); Acacia, Cajanus (Leguminosae); Nephelium (Sapindaceae); Lantana (Verbenaceae). Records for Pennisetum and Cajanus specified seed-feeding, and Bell noted flower feeding for the complex in India, as indicated above. The record from Durio is that of Piepers & Snellen (1904) from Java, so is most likely to be attributable to quadrimaculata.

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