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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.
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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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