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The concept
of the Nolidae has been expanded recently (Speidel et al., 1996;
Holloway, 1998; Kitching & Rawlins, 1998) to embrace the Sarrothripinae,
Chloephorinae and Camptolominae as well as the Nolinae sensu stricto as
treated here. The first two have been treated as noctuid subfamilies during most
of the last century (e.g. Hampson, 1912; Kitching, 1984). The third, though
associated with the other two by Mell (1943), has often been associated with the
Arctiidae (e.g. by Inoue et al., 1982); its status was reviewed by
Holloway (1988) in Part 6 of this series. The Nolinae have also often been
associated with the Arctiidae.
A detailed account of morphology within the group and a review of its
classification may be found in Holloway (1998), together with description of the
complex sound-producing (tymbal) structures found in males of many groups,
usually at the base of the abdomen, but also in single instances in the hindwing
(Cossedia Walker) and the valve, tegumen (Nycteola Hübner) and saccus of the genitalia (Cacyparis Walker). Sound production is also
encountered at the pupal stage, the pupa stridulating a row of beads or carinae
on the anterior margin of A10 against the interior of the cocoon, which may
itself be ridged, a phenomenon discussed in detail originally by Hinton (1948).
The monophyly of the group, discussed in the next section, and its separation
from the Noctuidae, increasingly seen as a paraphyletic assemblage (Kitching &
Rawlins, 1998; Mitchell, 1998; Holloway et al., 2001), is also being
confirmed in molecular studies, though on the basis of, as yet, a very small
sample of taxa (Mitchell, 1998; Mitchell et al., 2000).
Definition of the family
Holloway (1998)
noted seven characters, two of the pupal stage and five of the adult, that
define the family.
The most reliable feature, but one that is not known for any taxa that have not
been reared, is the structure of the cocoon: boat-shaped, with a vertical exit
slit, and a two-walled method of construction. The pupa lacks a cremaster, but
this is also a feature of the Euteliinae, currently in the Noctuidae. Some
probably plesiomorphic groups have the abdominal beading or carinae of the pupa
mentioned above.
In the course of dissection for this work, it became apparent that two of the
features noted by Holloway (1998) that provide support for the aedeagus in the
male genitalia are equally reliable and obviously more practical. The aedeagus
is supported by projections from the valves anteriorly into the abdomen:
dorsally there is a transtilla, extensions from the base of the valve costa that
fuse apically into a U- or V-shape (Speidel et al., 1996); ventrally,
similar processes extend from the base of each sacculus to fuse with a central
shield-like structure, referred to hereafter as the saccular shield. It is not
clear whether the central component of this second structure is homologous with
the juxta. The features can be appreciated particularly easily when dissecting
Risoba Moore species. The saccular shield can be seen
clearly in Figs 2, 9, 26, 168, 176, 180, 200, 212, 263, 308, 376, 394, 452, 456,
478, 497, 511, 522 and 542.
Speidel et al. (1996) noted a ventral displacement of the ventral part of
genital muscle m.4. Kitching & Rawlins (1998) considered that the elongate,
bar-shaped retinaculum of the male forewing, though shared with a few other
noctuoid groups such as the Arctiidae and Aganainae, was, when combined with
presence of scaling of the lower part of the clypeofrons, also diagnostic for
the Nolidae.
A final feature that needs further investigation is the observation by Holloway
(1998) that the patagia and tegulae easily become detached from the thorax when
this is compressed in the process of spreading specimens that have been relaxed
from a dry condition. The morphological basis of this is not clear.
Other features that are common in the family, but not necessarily unique to it
or found in all component groups include: the development of a pair of strong
lobes or apodemes on the basal margin of the male eighth tergite and sternite,
particularly the tergite (not in the Eariadinae); the presence of raised scales
on the forewing (mainly in the Nolinae, Sarrothripinae, Bleninae and
Collomeninae).
Several genera that have traditionally been associated with the groups now
incorporated in the Nolidae lack the defining features and should probably be
excluded (Holloway, 1998). These will be discussed in the final part of the
systematic account.
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).
Early stages and host-plant
specialisations
The larvae show a
great range of features, including proleg loss and development of secondary
setae on verrucae in the Nolinae as mentioned above. Much basic work on the
chaetotaxy and other features was done by Gardner (1941, 1946a, b, 1947, 1948a)
as reviewed by Holloway (1998) and Kitching & Rawlins (1998).
The bisetose/unisetose condition of the subventral setae on thoracic and
bisetose/trisetose condition of this group on abdominal segments are variable
within the family, but these features have been used to distinguish Noctuidae
from Arctiidae (T unisetose, A bisetose versus T bisetose, A trisetose
respectively). Thus Selepa and Earias show
arctiid features, and the Camptolomini have an arctiid thorax but a noctuid
abdomen. All other groups show the noctuid condition throughout. Other features
are discussed under individual groups.
Pupal characters and the method of cocoon construction have already been
discussed as defining the family; see also Gardner (1948b).
Some host-plant specialisation is seen in the family, or a non-exclusive focus
on a particular plant family. Both the Nolinae (Nola Leach) and the
Sarrothripini show a trend towards feeding on shoots, flowers, pods and seeds,
and one noline at least has been noted as a gall feeder (Itô & Hattori, 1982).
In the Careini, the Carea Walker complex of genera shows
a high level of feeding on Myrtaceae, and Aiteta Walker
is recorded only from the Combretaceae, particularly Terminalia.
Terminalia and Combretaceae are also the only hosts noted for Westermannia Hübner and
Miaromima
Meyrick in the Westermanniinae. Several records for Blenina
Walker and Labanda Walker are from Diospyros
(Ebenaceae), but these genera have also been noted from other families such as
Euphorbiaceae for the former and Dipterocarpaceae for the latter. Some
Collomeninae, such as Gadirtha Walker, appear to be
specialist on Euphorbiaceae. The Eariadinae show a temperate/tropical split
between Salicaceae and Malvales. Some of the ‘careine’ sequence of the
Ariolicini also appear to favour the Malvales.
Biogeography and ecology
The family is much more
diverse in the Old World, particularly the Oriental tropics. Only the Nolinae,
Sarrothripini of the Chloephorinae and Collomeninae extend to the New World,
although Baileya Grote has been associated with the
Risobini by Forbes (1954).
The matrices in Tables 1-3 provide biogeographic versus ecological profiles of
the major groups recognised in the Nolidae. Whilst the Ariolicini and Careini
show the high levels (around 40%) of endemics and Sundanian species with a
preference for lowland forest that are characteristic of a high proportion of
Bornean moth groups, the other two large groups, the Nolinae and Sarrothripini,
include many more widespread species that tend to have a higher ecological
amplitude, being found in both lowland and montane forests. The largest genera
of the Nolinae are Nola Leach and Manoba
Walker. Whilst the former shows the general character of the group, the latter
has a higher number of endemic taxa, most of which are montane, though this
endemic character may be an artifact due to the lack of data from other
mountainous parts of Sundaland.
Amongst the smaller groups, the
Collomeninae and the Westermanniinae show the more widespread type of profile,
whereas the Risobinae, in the form of the genus Risoba
Moore, have a greater proportion of endemics and lowland forest species, though
many of these fall within one lineage of the genus.
There are three large genera in the Careini, Calymera
Moore (21 species), Xenochroa Felder (36 species)
and Didigua Walker (19 species), that all show the
general Sundanian and endemic character of their tribe, though the first two
contain more montane species and the last, with over half its species restricted
to lowland forest, has a high proportion of those recorded from heath forest and
forests on acid soils generally.
Table 1. Percentage of species for Nolidae
groups, including the two largest genera in the Nolinae, amongst various
biogeographic and ecological categories as discussed in the text.
Table 2. Percentage of
species for Nolidae groups amongst various biogeographic and ecological
categories as discussed in text.
Table 3. Percentage of
species for large genera in the Careini and Risobinae amongst various
biogeographic and ecological categories as discussed in the text.
The sampling by Chey (1994) in softwood plantation and secondary forest habitats
indicates that the group as a whole maintains a moderate to high diversity in
these systems, particularly the Careini, Risobinae and Westermanniinae, and, to
a lesser extent, the Ariolicini and Sarrothripini. The ability of the group to
persist in cultivated areas is less clear, but probably not marked, except
possibly in some Sarrothripini and Selepa Moore. This
lack of persistence in more open and cultivated areas is probably a reflection
of the almost entirely arboreal larval feeding by the group.
Strong specialisation in larval host plant preferences is not common. However,
some groups appear to favour certain plant taxa, though rarely exclusively so.
Feeding on Fagaceae is frequent in the Chloephorini and Camptolomini,
particularly the more montane genera; montane Manoba species have also
been recorded from Fagaceae. Many careine genera show a high number of records
from the Myrtaceae, though Aiteta Walker species have
been recorded almost entirely from Terminalia in the Combretaceae, a host
also favoured by Westermannia Hübner and allies
in the Westermanniinae. Many of the host records for the more advanced
quadrifine Nolinae are also from the Combretaceae, and the family is
exploited by Risoba, along with the Melastomataceae and several other
families. Blenina Walker and Labanda
Walker are often recorded from Diospyros in the Ebenaceae, and several
genera in the Collomeninae appear to be associated with the Euphorbiaceae. The
Malvales are favoured by Gariga Walker and allies in the Chloephorini, Paracrama Moore and
Austrocarea
Holloway in the Ariolicini, and by the tropical Eariadinae generally.
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