SUBFAMILY EUTELIINAE
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Subfamily Characteristics
Adults

The species are generally more brightly coloured than most Noctuidae. The forewings are either rather elongate (Anigraea Walker, Anuga Guenēe) or squarish. All genera with squarish wings except Chlumetia Walker and Paectes Hubner have the margin angled more or less centrally, a character that appears to be more or less correlated with the presence of a subapical dark triangle based on the costa. The hindwing has a subtornal mark, often incorporated in a broad, dark border to pale ground colour. The hindwings below are usually finely multifasciate. Some wing venations are illustrated in Figs. 1-5.

Figures 1-5. Venation of Euteliinae. 1. Targalla palliatrix, 2. Penicillaria simplex, 3. Anuga constricta, 4. Anigraea mediifascia, 5. Paectes poliotis ().

Lateral scale tufts at the apex often give the male abdomen a squarish appearance.

The counter-tympanal hood is broader at the base than long, overhung
dorsally by a fan of short, rounded scales that arises from the metathorax above the tympanum; a tuft of slender scales arises anterior to the tympanum and extends posteriorly/obliquely to border the counter-tympanal hood ventrally.

The male antennae are characteristically broadly bipectinate over the basal half or two thirds, thence to the apex narrowly so or only ciliate (as below) in all Bornean genera except Anigraea where they are ciliate, Anuga where they are exceedingly long and narrowly bipectinate or ciliate in a taper towards the apex with no abrupt change, and Targalla Walker where they are filiform and smooth. The antennae of females are usually filiform though occasionally partially hipectinate as in the male. Partially bipectinate antennae of this form may be unique to the Euteliinae within the Noctuidae but are paralleled in other families such as the Cossidae, Limacodidae and Notodontidae.

The male genitalia are diverse in form, often strikingly modified, but the aedeagus vesica is usually basically spherical and the ductus ejaculatorius contains a globular sclerotisation that is also seen in the Stictopterinae. The eighth sternite in most genera bears a pair of eversible coremata within its basal half but in several genera (Anigraea, Atacira Swinhoe, Caedesa Walker, Chlumetia, Phalga Moore) it is modified distal to the coremata into a complex spined structure (Figs. 19-21, 24, 78-80, 104).


The female genitalia are more uniform within the subfamily. The bursa copulatrix is often ornamented with a pair of opposite signa centrally, each consisting of a coarsely scobinate indentation (Atacira, Chlumetia, Paectes). In other genera the bursa can be uniformly scobinate or with more distinctive ornamentation. In Eutelia Hübner the pair of signa are approximate rather than opposite, and in Kobestelia Gen. n. they are developed into interior blade-like processes.

The apophyses of the female eighth segment are broad and large in Phalga and Targallodes Holland, apparently modified into deep pouches in Eutelia, and reduced to small lateral processes in Chlumetia. In Aplotelia Warren they are present in diplographa Hampson, though very short, and have not been located in other species. In all the other genera they are absent, or possibly fused with the sclerotised basal portion of the ductus bursae either completely or sometimes still separate apically, appearing as lateral lugs subapically on the sclerotised part of the ductus.

The basal sternal sclerite of the abdomen is distinctive and may prove to define the subfamily. A basal, membraneous triangular zone is flanked by two interior, elongate lenticular flanges which extend distally from anterior apophyses and of which the plane is directed obliquely towards the centre of the sclerite (Figs. 17, 18). In all other noctuids examined for this character there is usually only a pair of grooves in a homologous position. The structure may serve to stiffen the sclerite and its development in the Euteliinae
from a groove (exteriorly; seen as a ridge interiorly) to a flange may be an adaptation associated with the characteristic resting posture of adults of the subfamily, described below.

The cryptic resting posture of adult euteliines is unusual, described by Bell in some of his manuscript life history accounts, e.g. for Anuga multiplicans Walker, and attributed by him to the subfamily generally. The wings are folded along lengthwise and the abdomen strongly curled up between them so that the wings appear like little sticks directed out horizontally, the forewings slightly forward, the hindwings often close to the abdomen. The resting insect thus resembles a dried up leaf or other plant detritus.

Larvae
Gardner (1948a) included the Euteliinae in his division C according to larval characters, particularly chaetotaxy. This division also included the Stictopterinae, Sarrothripinae but also taxa from several other subfamilies. The anterior prolegs are not reduced, or only slightly. The proleg crochets are usually homoideous (even in size), though heteroideous (irregular in size) in Anigraea, Anuga and Paectes. The setae are short, the spinneret moderately elongate, parallel-sided, and the spiracles are narrow.

Pupae
Gardner (1948b) defined the subfamily partially on the rounded apex to the pupal abdomen which lacks a cremaster (hook or spines used to anchor the pupa to the substrate or to a silken cocoon). But this character is also exhibited by the Chloephorinae and Sarrothripinae. The pupa of Phalga sinuosa Moore was noted by Gardner to have a circular patch of very regular, fine, longitudinal carinae at the posterior extremity; in all the other Euteliinae he examined this was smooth. In both Chloephorinae and Sarrothripinae there is often a bluntly spined or acutely corrugate ridge at the anterior edge of segment 10 ventrally.


HOST-PLANT RELATIONSHIPS AND ECONOMIC IMPORTANCE
Most of the species are defoliators but species of Chlumetia. bore in young stems and shoots of mango and are of economic importance.

Host-plant records have been garnered for this and the subsequent subfamilies from Gardner (1948a), Mathur (1942), Robinson (1975), Sevastopulo (1938-1947, 1941), Pholboon (1965), Browne (1968) and an unpublished manuscript in the BMNH by T.R.D. Bell (Bell, MS) and numerous records that have accumulated in a card index maintained by the Lepidoptera specialists of the Commonwealth Institute of Entomology.

There is a heavy bias within the subfamily towards plants of the family Anacardiaceae which includes the mango (Mangifera). A number of genera of the family are of economic use so many Euteliinae, particularly Penicillaria and Chlumetia species, have been noted as pests. Genera of Anacardiaceae from which euteliines have been reared in the Indo-Australian tropics are: Anacardium, Buchanania, Holigarnia, Lannea, Odina, Schinus, Semecarpus and Spondias as well as Mangifera. Other families recorded are Burseraceae (Bursera, Canarium, Garuga), Dipterocarpaceae (Anisoptera, Shorea), Leguminosae (Cajanus, Mucuna), Moraceae (Ficus, Morus), Myrtaceae (Eucalyptus, Eugenia, Myrtus, Syzygium; the Targalla delatrix group especially), Sapindaceae (Nephelium (litchi, rambutan, longan), Pometia), and Verbenaceae (Tectona).

In southern Africa (Pinhey 1975) Anacardiaceae are also recorded as host-plants (Pistacia, Rhus: Eutelia adulatrix Hübner) but also Leguminosae (Brachystegia, Baikiea: Targallodes polychondra Hampson), Myricaceae (Myrica: Eutelia bowkeri Felder) and Rutaceae (Citrus: Targallodes sub-rubens Mabille).

In North America (Tietze 1972) three species have been recorded from Rhus (Paectes oculatrix Guenee, Manathyssa basalis Walker and M. inficita Walker) but M. inficita includes Tsuga (Cupressaceae) and Ulmus (Ulmaceae) in its host range. The type species of Paectes, P. pygmaea Hübner, is recorded from Liquidambar (Hamamelidaceae) and P. burserae Dyar from Bursera (Burseraceae), the last family shared with the Oriental list.


ZOOGEOGRAPHY

The genera of Euteliinae present in Borneo exhibit a diverse range of geographical patterns.

Three genera, Penicillaria, Anigraea and Targalla, include a very high proportion of species that are found throughout the Indo-Australian tropics. Oriental/Australasian sister-pairs (Holloway 1982a) are found in Chlumetia, Paectes and Targalla. Chlumetia contains one widespread species, a group (with stout male genitalia) that are more localised and may replace each other allopatrically through the Indo-Australian tropics, a few localised Sundanian species, and possibly a montane group.

Both Targallodes vittalba Semper and Phalga sinuosa Moore are widespread Oriental species, the former extending to New Guinea. The allies of the Targallodes are African, but the Phalga has a sister species in Japan and Taiwan.

Caedesa and Kobestelia are purely Sundanian. Anuga and Atacira are virtually restricted to the Oriental tropics, both with centres of diversity in Sundaland.

Aplotelia and Paectes include a number of species complexes that range through the Indo-Australian tropics, the latter with a small group of species that have radiated within New Guinea. The wider complexes are treated in Figs 6 and 7.

Figure 6. Distribution of species of Aplotelia. 1.A. diplographa, 2.A. nubilosa, 3a. A tripartita tripartita, 3b.A. t.microsundana, 3c.A t.pratti, 4.A. oetakwa

Figure 7. Distribution of species of the cristatrix group of Paectes. 1. P.poliotis, 2. P.cristatrix, 3. P.psaliphora, 4. P.cyanodes, 5. P.languida, 6. P.fijiensis.

SYSTEMATIC ACCOUNT
All relevant generic type species (data in Nye 1975) were examined and both sexes dissected of each in order to establish the correct generic placement of the Bornean species. This has resulted in the generic names Eutelia Hubner and Phlegetonia Guenēe disappearing from the Bornean fauna; previously they occurred very frequently (Gaede 1937; Holloway 1976).

The genus Phlegetonia is based on the African species catephioides Guenēe. The male genitalia are of the same general form as those of the type species of Eutelia, the Mediterranean adulatrix Hübner. The genitalia of the females of these two species share several unusual characters such as approximation of the two signa, modification of the apophyses of the eighth segment to rounded pouches, and the presence of pockets of scales either side of the ostium bursae and laterally between segments 7 and 8 (four such pockets in all). Therefore Phlegetonia Guenee is here placed as a junior subjective synonym of Eutelia Hübner, syn. n.

The only true Eutelia species in S.E. Asia is E. adulatricoides Mell, widespread in tropical and subtropical Asia including Sumatra and Java but not Borneo.

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