The Lymantriidae are a major noctuoid family that is particularly
diverse in the Old World tropics. Current estimates of the number of species
worldwide range from 2160 (Holloway, Bradley & Carter, 1987) through 2500
(Ferguson, 1978; Common, 1990) to 2700 (Scoble, 1992). Heppner (1991) broke a
global estimate of 2416 down into regions that can be ranked as follows:
Afrotropical (1004); Oriental east to Moluccas (742); Australasia including New
Guinea and islands eastwards (255); Palaearctic (203); Neotropical (180);
The Australasian fauna has been estimated at 74 described species and
many undescribed by Nielsen, Edwards & Rangsi (1996). The lymantriid fauna
attenuates rapidly east of New Guinea with about 17 species in the Solomons
(Holloway, unpublished checklist), four in Fiji (an endemic trio of Calliteara
Butler, and the monobasic endemic Adetoneura
lentiginosa Collenette) and three in New Caledonia (of general Australasian
affinity, (see Orgyiini and Orvasca
Sciasticta Collenette comb.n., Orvasca
sp.2692, Orvasca rufalba
Holloway comb.n.). The family is absent from New Zealand.
Faunal richness decreases northwards from the Oriental tropics, with
just over 100 species recorded for Taiwan (Heppner & Inoue, 1992) and just
over 150 for Japan (Inoue et al, 1982). The Nearctic fauna (Ferguson,
1978) is essentially an attenuated Palaearctic one, though with the genus Dasychira
Hübner probably endemic (See Orgyiini). The Neotropical fauna is distinct from
that of the Nearctic and appears to bear no obvious relationships to that of the
Indo-Australian tropics (See Introduction).
The faunas of Borneo and Sumatra are similar in size and composition.
Schintlmeister (1994) referred to approximately 230 species for Sumatra, with
the comment that many more members of the Arctornis Germar (Redoa Walker)
group require investigation and description. He noted about 46 of these, a total
much lower than that for Borneo with about 81. Outside this group, the Sumatran
total of 180 is somewhat less than the Bornean tally of 216. The Bornean fauna
therefore contains over one third of the Oriental estimate of 742 by Heppner
Apart from the coverage by Schintlmeister (1994) of the Sumatran fauna,
adding greatly to the earlier treatment by van Eecke (1928), there have been
several other publications on island faunas in the region, particularly by
Collenette. These listed and described species in Peninsular Malaysia (Collenette,
1932), Java (Collenette, 1949a), Bali (Collenette, 1949b) and Sulawesi (Collenette,
1947), as well as being more broadly based geographically (e.g. Collenette,
1953). More recently there have been accounts of the faunas of Nepal (e.g.
Kishida, 1993, 1994, 1995) and Taiwan (Wang, 1993).
The highly polyphagous, but largely arboreal larvae include many
agricultural and forestry pests and, when abundant, may also cause medical
problems with the urticating properties of their setae (see Nygmiini
n and Toxoproctis
Gen.n.). On the
other hand, their richness in tropical lowland forest in the region and
relatively low mobility may enable them to be used as indicators for
biodiversity assessment and environmental monitoring (Holloway, 1984, 1985b).
Definition of the family
The Lymantriidae are noctuoids with a thoracic tympanum and show a number of
unique features within the superfamily (Ferguson, 1978; Maes, 1984a; Common,
1990; Scoble, 1992), as well as having a prespiracular counter- tympanal hood as
in the Arctiidae.
The tongue is usually reduced or absent: the adults do not feed.
The antennae of both sexes are strongly bipectinate, the pectinations being
longer in the male, terminating in one to three long spinules (Figs 1, 2),
though such spinules also occur in some Arctiinae (Kitching & Rawlins, 1999
Males often have paired, pocket-like tymbal organs on the third abdominal
sternite (Figs 3-7). The occurrence of this character was reviewed by Dall' Asta
(1988). It is found in many genera of all the tribes occurring in Borneo and in
all the genera not placed to tribe except one. It may therefore be part of the
family ground-plan. However, it was only found in one (Thagona Möschler)
of a sample of four Neotropical genera dissected.
The female genitalia have a small pair of setose lobes ventral to, and distinct
from, the ovipositor lobes. These were termed pseudopapillae by Maes (1984a),
who suggested they were diagnostic for the family, but they are seen also in
aganaine Noctuidae (Holloway, 1988) and some Arctiidae (Kitching & Rawlins,
1999 ). They can be seen, for example, in Figs 81, 214 and 243.
The larvae have conspicuous, round, red or yellow dorsal glands, one on each of
abdominal segments 6 and 7.
The adult resting posture is tectiform, or flattened tectiform, with the hairy
front and middle legs displayed, the forelegs held out in front, the middle legs
laterally (Tweedie & Emmet, 1991). However, some lithosiine arctiids exhibit
a similar resting posture (Kitching & Rawlins, 1999 ).
observations are drawn from accounts of the family cited in the previous
section. The adults show a great diversity of wing facies, though it is usually
only the forewing that is patterned. This can range from the rather typical
noctuoid patterning of most Orgyiini, through the more zig-zag fasciation of the
Lymantriini, to the bolder, more colourful patterns of the new tribe Nygmiini or
the satiny white of most species in the new tribes Leucomini and Arctornithini.
Ocelli and chaetosemata are absent. There are often long, hair-like scales on
the thorax and abdomen. Both forewings and hindwings have M2 arising closer to
M3 than to M1, the quadrifid state of the former being common to all noctuoids
except the Notodontidae. A forewing areole can be present or absent.
Sexual dimorphism is moderate to extreme, females tending to be much larger,
particularly in the abdomen which, with the non-feeding habit, is full of eggs
at eclosion. Females either have much larger wings than males or, in the case of
some Orgyiini, are flightless and brachypterous or apterous. Even when winged,
most lymantriid females fly only weakly and are much rarer than males in
Most female lymantriids have a scale tuft (corethrogyne) at the apex of the
abdomen, and this is used to cover the egg mass to protect it. The scales can be
irritant. Stereoscan photographs of these scales from egg masses of a small
sample of lymantriid species made at the Australian National Insect Collection
suggest that these may vary amongst the tribes. Those of Lymantria are
narrow, long, longitudinally ridged, the ridges finely serrate such that they
are barbed away from the apex. Between the ridges are frequent septa, with some
perforations between the septa. The scales of Orgyia are similar, but
mostly broader and of more variable breadth. In the Nygmiini they vary from
ridged to irregularly and strongly barbed rods, the barbs of variable length and
frequency, more like thorns on a stem: the latter could be larval hairs derived
from the cocoon, as described by Kitching & Rawlins (1999 ). Weakly
flighted or apterous females may lay their egg mass on the cocoon from which
they have just emerged. The extent to which this occurs is variable but is most
developed in the Nygmiini and Orgyia. The Arctornithini are exceptional
in laying eggs singly, the females lacking a corethrogyne. The Leucomini are
intermediate, laying eggs in rows or groups, lacking a corethrogyne, but with
scales attached to the egg mass in Leucoma (de Worms, 1979).
The male genitalia are highly variable in structure. The eighth abdominal
sternite is rarely modified to a large extent (an exception being Dura Moore),
though the tergite is occasionally so, as in Dura, Calliteara of the
Orgyiini, some genera of the Nygmiini and in one of the unassigned genera. The
female genitalia usually have distinctly setose ovipositor lobes, short and
deep, as well as the pseudopapillae. The signum is bicornute or a flange in all
tribes except the Arctornithini (see below and Arctornithini n). The development of the
sterigma is highly variable, but reaches its greatest complexity in some
The eggs are of the upright type, circular, domed or slightly flattened, laid
usually in tight masses, though the Arctornithini and Leucomini are exceptions.
The larvae can remain gregarious (e.g. some Nygmiini) or disperse, often by
ballooning on silken threads (Orgyiini in particular, where the female is
flightless or only weakly so).
The larvae have dense and often long secondary setae on verrucae as in other 'hairy' noctuoids such as Arctiidae. The urticating properties of these have been
referred to. Denser brushes or longer hair pencils occur in some groups.
Crochets are numerous on the prolegs, in a uniordinal, homoideous mesoseries.
The disposition and size of the verrucae and other larval characteristics will
probably prove of value in elucidating relationships within the family, as
indicated by the key of Gardner (1938) and discussed in later sections. Gardner
presented a very detailed account of Indian species from all five of the tribes
Pupation is usually in a dense silken cocoon that incorporates larval setae,
though this can be very weak or vestigial in the Leucomini and Arctornithini
where the pupa is exposed in a cradle of silk. The pupa also has dense secondary
setae, mostly arising from the verrucal scars of the larva. A cremaster is
present, with short, hooked setae at its apex.
Classification within the family
(1978) considered the possibility of segregating subfamilies within the
Lymantriidae but felt that knowledge of tropical faunas was too limited to
permit this. He contented himself with segregating Nearctic taxa into two
tribes, Lymantriini and Orgyiini.
Though this Bornean survey has probably covered most of the morphological
variety present in Indo-Australian representatives of the family, there are
sections of the Afrotropical fauna that require further study and the whole of
the small Neotropical fauna. Dissection of a sample of Neotropical genera (Caviria
Walker, Desmoloma Felder, Sarsina Walker and Thagona Möschler)
has shown no definite relationships with any of the groupings or isolated genera
recognised for Borneo. Only Thagona males had tymbal organs and none of
the females examined had a signum. A number of Neotropical genera thought to
have been Lymantriidae are now placed in the Bombycidae as subfamily Phiditiinae
(Lemaire & Minet, 1999 ).
Therefore, it was decided to continue the process started by Ferguson and to
identify tribal groupings rather than subfamilies. The result is the recognition
of three new tribes, segregated from Ferguson's concept of the Lymantriini. They
are discussed in more detail in the relevant parts of the systematic section,
but are listed below with their principal diagnostic features.
forewing facies is characteristic, with zig-zag or lunulate fasciation,
a V-shaped discal spot and an orbicular spot.
larva has dorsal brushes on the first four abdominal segments.
seventh abdominal segment is expanded and membranous in the female.
male genitalia are asymmetric.
signum of the female bursa is a broad, sparely but evenly scobinate
addition, there are six genera treated at the end that could not be assigned
with confidence to any of these groups.
Relationships within the Noctuoidea
Kitching & Rawlins (1999 ) have suggested that the
Lymantriidae are sister-group to a paraphyletic Pantheidae, sharing apomorphic
features such as the presence of secondary setae in first instar larvae,
including, on the prothorax, distinctive, enlarged prespiracular verrucae,
making that segment the widest part of the body. The character is otherwise seen
only in some Acronicta.
As both the Pantheidae and Acronicta have already been covered in
this series, the former as a subfamily of the Noctuidae (Holloway, 1985a), and
the latter within a narrow concept of the noctuid subfamily Acronictinae
(Holloway, 1989a), a brief review of their classificatory position is merited.
There is widespread disagreement on this position. Poole (1995) placed the
pantheines and acronictines in a group with the noctuid subfamily Bryophilinae
on the grounds of the presence of secondary setae in first instars, a character
he interpreted as a reversal to a primitive state. Otherwise, he stated that the
adults of the three groups are ‘clearly “trifid” noctuids on the basis of
larval characters and the structure of the tympanal region’, though later he asserted that ‘I have an unsupported feeling that the Pantheinae and
Acronictinae are not closely related despite the similarity of their larvae’.
Speidel, Fänger & Naumann (1996) also treated the pantheines as a noctuid
subfamily, grouping them in a trichotomy with the Nolidae (as defined by
Kitching & Rawlins (1999 ) and by Holloway (1998a)) and a large clade
that includes the trifine subfamilies covered by Holloway (1989a) with the
Plusiinae and a few other small groups. There is an emerging consensus on the
broad composition of this clade, though disagreement over the membership of some
of the groups, particularly the Pantheinae (idae), though all place the
Acronictinae sensu stricto within it.
It is likely that the Pantheinae do not represent a monophyletic group, and
there is disagreement over which genera should be included. Speidel, Fänger
& Naumann (1996) included Dilobinae and Mominae in their concept: neither of
these groups is represented in Borneo. Kitching & Rawlins (1999 )
included all the Bornean genera in their Pantheidae, but placed Dilobinae in the
The Bornean pantheine species covered by Holloway (1985a) consist of three
rather dissimilar species in different genera, all with quadrifine hindwing
venation and conspicuously hairy eyes. Trichosea Grote may be related to Acronicta,
as discussed by Holloway. Elydnodes Hampson has genitalic features in
common with Trichosea and might best be grouped with that genus, wherever
it is finally placed.
The other two species, in the genera Anepholcia Prout & Talbot and Antitrisuloides
Holloway, together with Trisuloides Butler and Smilepholcia Prout
& Talbot, belong to a predominantly montane Indo-Australian assemblage of
rather large species with densely scaled, variegated forewings and orbicular
stigmata. The hindwings are yellow, bordered broadly with black interior to
yellow fringes. The relationships of this group require further investigation,
particularly with regard to the north temperate pantheine type genus, Panthea
Hübner. This will probably require discovery of the early stages. Kobes
(1985) has covered the slightly more diverse Sumatran pantheine fauna.
In conclusion, the precise relationships of the Lymantriidae within the
Noctuoidea are still unclear, but they may group with the Arctiidae, Nolidae and
possibly part of the Pantheidae in a sister-relationship with the bulk of the
groups currently placed in the Noctuidae (Kitching & Rawlins, 1999 ).
Recent molecular studies (Mitchell, 1998) strongly support a sister-relationship
between Lymantriidae and Arctiidae, but place these as sister to another pair of
taxa currently treated as Noctuidae subfamilies by Kitching & Rawlins (Aganainae
and Herminiinae) but by other authors (e.g. Common, 1990) as good families. The
molecular data also placed the Pantheinae with the trifine Noctuidae in a
The Arctiidae, Nolidae (Holloway, 1998) and Lymantriidae all have sound-
producing organs in all or a large number of their taxa, though the location of
these is different in each case. Kitching & Rawlins suggested that these
features might be serially homologous (sternal) and therefore provide some
evidence of relationship. Sound production in Noctuidae is mostly restricted to
trifine groups (Heliothinae, Agaristinae), where it involves foveate areas of
the forewings (Matthews, 1991); foveae also occur at the base of the forewings
of Amyna Guenée species (Acontiinae). Amongst the Catocalinae, Arcte Kollar
has a conspicuous strigil on the sixth abdominal tergite (Robinson, 1975).
As stated earlier, lymantriid larvae are, as a rule, highly polyphagous,
and this is general for all the tribes recognised here. The larvae are
predominantly arboreal defoliators, but a few orgyiines and nygmiines also
attack herbaceous plants. In the former, the genera Psalis Hübner, Laelia
Stephens and Pantana Walker have become specialist on grasses and
reeds, the facies of the adults also reflecting this in its rather straw-like
The other instance of specialisation is by the leucomine genus Perina
Walker that has, apart from a few dubious records, been recorded entirely
from Moraceae, particularly the banyan, Ficus religiosa.
Some lineages of Nygmia Hübner may favour Loranthaceae, and the
diversity of Arctornis Germar in Sundaland may reflect an ability to cope
with the challenges of dipterocarp foliage rarely seen elsewhere in the
Lepidoptera (See Arctornithini n).
Ecological and biogeographic representation
Table 1 presents summaries of the biogeographic and ecological
composition of the four largest tribes in Borneo in the form of two-way tables,
biogeographic categories against ecological ones. This format has been applied
in previous volumes of the series and originally by Holloway & Barlow (1992)
and Holloway (1994).
Well over half the species in all tribes except Orgyiini fall into the Sundaland
or endemic lowland forest categories. Endemism is highest in the Arctornithini,
though this may decline as the faunas of other parts of Sundaland become better
known. It is also substantial in the Nygmiini but low in the Lymantriini. The
Orgyiini have more species in the widespread biogeographic categories and the
disturbed, open habitat category, a reflection of the high level of pest species
in the group and the adaptation of many to feeding on herbaceous vegetation.
Montane species are infrequent except in Orgyiini, where these are again of a
more widespread nature, and Nygmiini, where endemism is high. All four groups,
but particularly the Nygmiini and Arctornithini, are well represented in the
ecological category for high ecological amplitude over altitude, with many
species occurring in both lowland and montane forest.
The family generally is extremely rich in primary lowland forest and declines in
diversity with latitude, altitude and disturbance. Chao (1981) demonstrated a
similar decline with altitude on Mt. Emei in western China, with 93% of species
recorded below 1100m and only 21% above 1900m. Three- quarters of this fauna is
Oriental tropical and subtropical, the rest Palaearctic, with the latter
increasing in representation with altitude.
Holloway (1984) noted a gradient in lymantriid diversity in forest types sampled
during his survey of the G. Mulu National Park, with diversity increasing from
young secondary forest through alluvial forest regenerated from farmland
abandoned a century before, to undisturbed lowland forests. Data for secondary
forest and various types of softwood plantation in Sabah were obtained by Chey
(1994). The general trends of moth diversity observed in these sites (Chey,
Holloway & Speight, 1997) are also evident in just the Lymantriidae, with
the samples from secondary forest, Eucalyptus deglupta plantation (where
there was a much more diverse secondary forest component in the understorey) and
one of Pinus caribaea having higher diversity than the others, the lowest
being from the two legume tree plantations (Acacia mangium, Paraserianthes
falcataria), where high numbers of Orgyia postica Walker and, in the
former only, Rhypotoses glebula Swinhoe were recorded. Values of the
Fisher, Corbet & Williams (1943) a measure for a selection of these sites
are given in Table 2. The Mulu survey values and those from the data of Chey are
not strictly comparable, as the former were from pooling two to four consecutive
nights of samples and the latter from pooling samples made over a whole year.
Mean values of a for pooled monthly samples from the plantations were half (or
less) those for the same sites for the whole year for all Macrolepidoptera
(Holloway, 1998b). The gradient of diversity appears to be much stronger in the
Arctornithini than in the Nygmiini.
Table 1. Percentage of species for the major tribes of Bornean
Lymantriidae amongst various biogeographic and ecological categories as
discussed in the text.
Measures of lymantriid diversity from light-trap samples made in lowland forest
types during the survey of the G. Mulu National Park, Sarawak (Holloway, 1984)
and by Chey (1994) in softwood plantations near Brumas in Sabah.
||α (95% confidence)
Forest on limestone
Heath forest on river terrace
Hill dipterocarp at 500m
Hill dipterocarp/alluvial boundary (150m)
Alluvial forest (regenerated)
||23.5 (± 4.9)
||21.7 (± 5.2)
||16.4 (± 3.8)
||14.2 (± 2.4)
||12.1 (± 2.0)