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Groupings within the family
A tentative classification into
subfamilies and tribes is beginning to emerge (Holloway, 1998; Kitching &
Rawlins, 1998; Holloway et al., 2001), but there is great disparity in
the size of the divisions, some, such as the Nolinae and Chloephorinae, being
very large, and others being monogeneric, such as the Bleninae, Risobinae
(excluding Baileya Grote) and Eariadinae. There are also
several genera and small generic groups that do not fall readily into these
categories but nevertheless show typical nolid features.
The Nolinae (see also Holloway & Miller (1995)) are defined essentially on
larval features such as reduction and loss of the first pair of prolegs, and
presence of secondary setae on verrucae. The adults lack ocelli; these are
present in the rest of the family and in Beana Walker, possibly the most
plesiomorphic noline. There are also features of the male and female genitalia
common to many genera.
The Chloephorinae assemble a number of tribes where at least some genera have
tymbal organs at the base of the male abdomen. These show their greatest
development and ubiquity in the Chloephorini (Figs 208, 211, 212, 214, 215) and
Camptolomini, but also occur in all Careini (Figs 228, 230-239). In the
Ariolicini (Figs 360-364, 396) and the Sarrothripini (Figs 126-131) they are
present in only a few genera, though others have structures on the male basal
sternite that may represent vestiges. These last two tribes are the most weakly
defined. The Ariolicini have two subgroups, genera of the first mostly having
peg-like setae on the valves or strong hair pencils at the valve bases, and
those of the second having male genitalia bearing a possibly superficial
resemblance to those of some Careini. The Sarrothripini frequently have a
process on the valve that bears dark scales or setae, though this is not seen in
the type genus, Nycteola Hübner. They usually have rather grey forewings
with raised scales. Females of many genera have acute ovipositor lobes. These
two tribes are the only ones in the Chloephorinae where pupae with beading have
been noted.
No strong signals of relationships have been located in the head, wing venation
or female genitalia (Holloway, 1998). The male antennae are generally
fasciculate (filiform, with cilia), though bipectinate ones are seen in many
Nolinae, two species of Labanda Walker in the Ariolicini and the
Gelastocera Butler
generic group. The absence of ocelli in Nolinae has been mentioned earlier. The
labial palps generally have the third segment much shorter than the second,
exceptions being discussed under
Cacyparis Walker, which, with
Ballatha Walker, has the third segment very long and slender. Reduction of
the tongue is seen in several genera of the Ariolicini and some Collomeninae.
The groundplan venation is typical of the collection of groups currently
assigned to the quadrifine Noctuidae subfamilies. The forewing has an areole,
often elongate, at the anterior angle of the cell; this areole gives rise to the
radial sector veins, usually with R2 and R5 arising independently from it on
either side of the stalked (R3, R4). The hindwing typifies the quadrifine
condition, with M2, M3, CuA1 and CuA2 arising independently, but often with M3
and CuA1 connate, around the posterior angle of the cell.
In a number of nolid groups this groundplan becomes modified. Stalking and
reduction of hindwing veins is seen in the Nolinae, Sarrothripini, Ariolicini
(loss only in
Titulcia Walker, but an unusual branching system in
Chandica Moore) and
Eariadinae, and to a lesser extent (occasional stalking of M3 and CuA1) in the
Chloephorini and Careini; in
Phaeothripa Hampson and
Ptisciana Walker
of the Collomeninae M2 and M3 are stalked. Variations on the forewing groundplan
consist of reduction and loss of the areole, and variations on the radial sector
branching pattern, for example, with R5 branching from Rs more distally than R2
in some Sarrothripini and independently from the cell in others. In the
Eariadinae and Chandica and
Cossedia Walker in the
Ariolicini R2 is independent but not R5. Radial sector veins are lost in some
Nolinae such as Nola
Leach.
The male genitalia show the aedeagus support structures that define the family.
Reduction of the uncus is common, and there are often scaphial thickenings on
the anal tube. The tegumen may be elongate, ventrally swollen on each side,
extending beyond the junction with the vinculum and often bearing hair pencils (culcita
of Kobes (1997)). The Chloephorinae and some other groups frequently have a
subbasal process on the valve costa. A saccular harpe or other structure is less
common, but is seen in the Nolinae, Risobinae, Bleninae, Westermanniinae and the
genera Selepa
Moore and
Bryophilopsis Hampson. The valve is often paddle-shaped, with basally
directed hair-setae distally on the paddle, which may be bilobed; these features
are seen particularly in the Careini and the second group of the Ariolicini.
In the female genitalia the ovipositor lobes can vary from rounded, ring-like
(many Ariolicini) to acute (many Sarrothripini). The sterigma is not often
modified, and the ductus bursae is very variable in length. The occurrence and
form of the signum or signa is also diverse, and general scobination of the
bursa is also frequent.
In four widely separated genera (Mniothripa
Hampson, Tympanistes
Moore, Westermannia
Hübner, Didiguides
Kobes) the cornuti on the aedeagus vesica are deciduous, become detached during
copulation and can be found in the corpus bursae of a mated female (Figs 192,
286, 299, 484). In all these cases the cornuti are somewhat dagger-like. This
phenomenon is not unique to Nolidae but also occurs in, for example, ennomine
Geometridae (Holloway, 1979: 338) and Notodontidae (Holloway, 1983: 4).
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