The Notodontidae have a metathoracic tympanal organ and are therefore in the superfamily Noctuoidea (Brock 1971) or placed in a separate superfamily, Notodontoidea, associated with the Noctuoidea (Common 1970, 1975). The former arrangement is perhaps preferable as the thoracic tympanal organ is one of very few really reliable apomorphic (derived rather than primitive) characters in the higher classification of the Ditrysian Lepidoptera.

Characters for defining the Notodontidae themselves are less satisfactory but there is general agreement that they retain a higher proportion of primitive, or plesiomorphic, characters than other families in the Noctuoidea (e.g. indications of median veins in the wing cells of Phalera bucephala Linnaeus), and hence may be the sister group to the rest. Both Brock (1971) and Common (1970) cited the possession of a metascutal tympanal bulla as a character defining the family, and Common also regarded the ventral direction of the tympanum itself as diagnostic, though this character is not infallible. In his key to superfamilies Common (1970) distinguished his Notodontoidea from the Noctuoidea using a character of wing venation: the base of vein 5 (M
2) is usually nearer vein 6 (at the anterior angle of the end of the cell) than to vein 4 (at the posterior angle of the cell) rather than vice versa. Again this is not infallible.

Several characteristics of the male genitalia and abdomen may prove to be of value either in defining the family or in subdividing it.

Many species have lateral spurs to the fourth abdominal sternite that bear a fringe of flattened, blade-like setae
*; the fourth sternite is also often distally bilobed, the lobes bearing more persistent scales or fine hairs marginally. This character is seen, for example, in Ambadra Moore, Caschara Walker (Fig. 90), Archigargetta Kiriakoff, Phalera Hübner, Brykia Gaede, Ginshachia sumatrensis Gaede (but not G. bronacha Schaus), Allata Walker and Saliocleta Walker (e.g. barasamphia Schaus).

The majority of genera have some sort of modification to the male eighth sternite, sometimes also the tergite, but there is great variety in these modifications and they cannot be categorised in terms of a typical 'notodontid form' though they may prove of value in subdividing the family or recognising groupings of genera.

The presence of deciduous stellate spicules in the aedeagus vesica is another character widely distributed in the family, e.g. in Phalera, Hyperaeschrella Strand, Brykia, Epistauropus Gaede, Euhampsonia Dyar, Teleclita Turner, Pseudoteleclita Kiriakoff, Disparia Nagano, Oxoia Kiriakoff, Notodonta Ochsenheimer, Suzukiana Sugi, Allodonta Staudinger and Hexafrenum Matsumura. In some instances the stellate spicules become asymmetric with one spine of each spicule being emphasised, the other spines fringing the base (when still attached to the vesica) as in Turnaca punctata sp. n. and, reduced to one massive spine at the end of a long, tube-like vesica, in Caschara punctifera Walker. In Ambadra and Fentonia Butler there are merely many simple, small, needle-like deciduous spicules. These can be of value in assigning males to females in certain species as a female that has been mated will retain them in her bursa copulatrix, and their size and shape may be diagnostic.

Almost all species have the gnathal processes separate, robust, prominent, and often complex. The uncus itself is usually modified to some degree, sometimes with elaborate socii.

A large number of species have the saccular area of the valve of the male genitalia membranous and somewhat corrugate, another character probably unique to the Notodontidae in the Noctuoidea. This character is seen in species of Euhampsonia, Calyptronotum Roepke, Chadisra Walker, Caschara, Allodonta, Hexafrenum, Hyperaeschrella, Disparia, Notodonta, Clostera Samouelle, Micromelalopha Nagano and Plusiogramma Hampson.

The forewing pattern is generally noctuoid with reniform and orbicular stigmata and simple postmedial and antemedial fasciae. The hindwings are usually unmarked except sometimes along the costa (Quadricalcarifera Strand and relatives, Stauropus Germar and relatives) or at the tornus (the Chadisra Walker and Teleclita Turner groups of genera). The presence of such hindwing markings may not be of great taxonomic significance but merely indicative of exposure of that area of the hindwing when the moth is at rest. As mentioned in the Introduction, the forewing patterns are cryptic, suggestive of tree bark or lichen; species with lichenous or bark-like patterns usually rest with the wings obtusely angled over the body. Other species, such as in the genus Phalera Hübner or in the Gargetta Walker group, rest with the wings scrolled round the body, causing them to resemble broken twigs.

In many species the dorsum of the forewing bears a central angular projection. Examples are Norraca lativitta Walker and the genera Suzukiana, Hyperaeschrella, Peridea Stephens, Notodonta, Hexafrenum, Caschara, Allata and allies, and Rodneya Kiriakoff.

The forewings and abdomen are usually long and narrow, the abdomen extending well beyond the hindwing margin. The antennae are usually bipectinate or pectinate in the male, but sometimes filiform with ciliae. The female antenna is filiform in the majority of cases, less elaborate than the corresponding male antenna.

Gardner (1943) stated that the larval characteristics justified inclusion of the family in the Noctuoidea and drew attention to the resemblance of those notodontid larvae with normal prolegs to the larvae in the noctuid subfamily Acronictinae.

All four pairs of prolegs are present and nearly equal except in Gargetta Walker and allies, where the first, and sometimes the second, are reduced. The claspers have crochets in a uniordinal homoideous mesoseries (a single uniform band) and always with conspicuous secondary setae (Gardner 1943).

The generic characters are sharply definable and will no doubt lead to much revision of the current taxonomy when more life histories are known; they may even provide a basis for a rational subdivision of the family. Features include a range of form of the anal prolegs (or claspers) from vestigial through normal to slender and stematopodiform.

Stematopodiform claspers are often held out horizontally or erect and may contain protrusible lashes. They are seen in the stout, anteriorly squarish larvae of the Cerura Schrank group of genera and in the slender, cylindrical larvae of the Gargetta group of genera. The Australian species Pheressaces cycnoptera Lower (Turner 1903) and the Indian Fentonia tenebrosa Walker (unpublished notes by T.R.D. Bell) also have a Gargetta-like larva and the Indian species must therefore be misplaced in Fentonia Butler.

Another, perhaps natural group of taxa with modified anal prolegs has these filiform, vestigial, set on dorsoventrally flattened terminal abdominal segments, the latter resembling a leaf to some extent. The anterior part of the larva is also modified, usually by a number of dorsal humps. The development is at its most extreme in larvae of Stauropus Germar and Neostauropus Kiriakoff, but is also seen, with the leaf-like appearance most definite, in Teleclita and Pseudoteleclita and their African relatives in the genera Afroplitis Kiriakoff and Amyops Karsch (Pinhey 1975).

Many other genera have dorsal prominences of some sort and most are strikingly, if cryptically, patterned.

The literature on the early stages of Oriental tropical and Australasian species is sparse and scattered but an attempt is made in the main text to collate and precis what there is and assemble all the host-plant data. A major source of information was an unpublished typescript by T.R.D. Bell in the British Museum (Natural History) (BMNH), referred to in the text following as (Bell, MS), accompanied by watercolours of many of the species described, reared in South India. The adult material reared was deposited in the BMNH so it has been possible to check the identity of the species. Other sources are Barlow (1982), Bell (1935), Dodd (1902), Gardner (1943, 1946), Issiki (1969), Kalshoven (1981), Pant & Chatterjee (1951), Sevastopulo (1938-1947, 1940, 1949), Sugi & Nakatomi (1969) and Turner (1903).

The cteniophores of Jordan (1923) who suggested they were associated with abdominal glands and and scent scales on the hindwings or legs.


The references listed at the end of the discussion on larvae, together with Browne (1968), Pholboon (1965) and unpublished records in a card index maintained by Lepidoptera specialists of the Commonwealth Institute of Entomology, are the sources used in this survey of host-plant relationships and in the discussions of individual species later.

Several interesting patterns or relationships are becoming evident despite the sparseness of the data.

The majority of species defoliate dicotyledonous trees and shrubs but the Pydna Walker and Ambadra Moore groups of genera, together with Phalera combusta Walker (extending from India to Sumatra but not Borneo), feed on monocotyledons, mainly palms, bamboos and grasses. The streaky fawn, yellow or red forewing patterns of these groups reflect their feeding habit, paralleling the adults of other monocot-feeding species in other families such as the Mythimna and Sesamia groups in the Noctuidae, the crambine Pyralidae, and Psalis pennatula Fabricius in the Lymantriidae.

Several species are polyphagous over a range of plant families, such as Neostauropus alternus Walker and the Chadisra species from India. Species of the genus Phalera tend to be polyphagous but Oriental tropical species are found more frequently or exclusively (P. javana Moore) on Leguminosae.

A preliminary survey of host-plant relationships in the Indo-Australian macrolepidoptera has pinpointed a number of genera in other families that are restricted to Leguminosae such as, in the Ophiderinae (Noctuidae), the genera Lacera Guenee, Ericeia Walker and Rhesala Walker. The Notodontidae probably also include legume specialists, most conclusively the Allata Walker group of genera - Allata, Eguria Matsumura, Rosama Walker (Bell MS; Gardner 1943, 1946). Besida xylinata Walker has also been recorded from the family (Pholboon 1965).

Netria viridescens
Walker would appear to be restricted to the Sapotaceae, feeding on several genera of the family such as Mimusops, Madhuca, Sideroxylon (Bell MS) and Manilkara (CIE records) in India.

The genus Dudusa Walker may prove to be restricted to the Sapindaceae, with D. synopla Swinhoe recorded from Schleichera in India (Bell 1935, as nobilis Walker), a record from Thailand on Nephelium also attributed to nobilis (Pholboon 1965) and D. vethi Snellen recorded from Nephelium in Sumatra and Java (Kalshoven 1981, as nobilis).

The genera Teleclita and Pseudoteleclita have only been recorded from Terminalia (Combretaceae), in India (Bell MS), the Philippines (original material of T. cathana Schaus) and Australia (Dodd 1902; Turner 1903). African relatives in Afroplitis, Amyops and Galona Karsch have similar larvae that feed on Terminalia and Combretum in the Combretaceae but also on Parinari in the Chrysobalanaceae and guava (Myrtaceae) (Pinhey 1975). Two chloephorine noctuid genera, Aiteta Walker and Westermannia Hiibner, also appear to be restricted to Terminalia in the Indo-Australian tropics but this plant genus is host to an unusually large number of Lepidoptera species, mostly polyphagous ones.

An interesting parallel in host-plant relationships is evident between the Clostera group of genera and the Cerura group; it is also shown by the nymphalid butterfly Phalanta phalantha Drury (Corbet & Pendlebury 1978). In the Palaearctic Region the host plants are Salicaceae, especially Salix and Populus, but in the tropics the majority of hosts are in the Flacourtiaceae, genera such as Casearia, Flacourtia, Scolopia (in Australia (Common 1963)) and Xylosma being recorded. In India both plant families have been noted as hosts and also in Hong-Kong (M. Bascombe in litt.). Terminalia is also involved in India (Brown 1968). Notocerura spiritalis Distant, an African member of the Cerura group with a characteristic larva, also has Flacourtia as a host (Pinhey 1975).

This relationship is less exclusive in Clostera than in the Cerura group. Records from a few other plant families have been made for the two major species groups in Clostera that extend into the Palaearctic. In the anachoreta-fulgurita group Palaearctic records are from Salicaceae. In India they are from both families (Browne 1968; Gardner 1943; CIE records) and also from Elaeodendron in the Celastraceae (Gardner 1943). There are no records from South-east Asia but the group has been reared from Flacourtiaceae in Hong-Kong (M. Bascombe in litt.). The anastomosis-restitura group is known from Salicaceae in the Palaearctic and Hong-Kong, from both families in India but also from Elaeocarpus (Elaeocarpaceae) there (Sevastopulo 1940). The Melanesian representative of the group, rubida Druce, has been reared from Terminalia in Papua New Guinea (CIE records). Southern African Clostera feed on Proteaceae (Pinhey 1975).

The two plant families are not taxonomically related according to Willis (1973), the Flacourtiaceae being linked with families such as the Passifloraceae, Euphorbiaceae and Tiliaceae rather than the Salicaceae.

There are several host-plant records for the Gargetta group of genera (Bell MS; Gardner 1943; Bascombe in litt.). Porsica ferreopicta Hampson, P.ingens Walker and Phycidopsis albovittata Hampson have all been recorded from Antidesma, Gargetta costigera Walker and G. ?divisa Gaede from Briedelia, Porsica punctifascia Hampson from Aporusa and Porsica curvaria Walker from Bischofia. All four plant genera were previously placed in the Euphorbiaceae but only Briedelia and Aporusa are still included (Willis 1973). Antidesma is the only genus of the Stilaginaceae, suggested by Willis to be intermediate between Icacinaceae and Euphorbiaceae, and Bischofia is likewise unique to its family, the Bischofiaceae, related by Willis to the Staphyleaceae with a comment that relationship with the Euphorbiaceae was probably illusory. The association of the three plant genera in respect of their utilisation by what is probably a natural group of Lepidoptera might indicate that their taxonomic relationships should be reexamined.

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