The information on
distribution and habitat preference is summarised in the following table and in
+ New Guinea
+ N.E. Himalaya
(+ Wallacea) (+ Taiwan)
tropics to Sundaland
tropics to Wallacea
tropics to New Guinea
Shared with Sumatra only.
Shared with Peninsular Malaysia only.
two species in the ‘Other’ category are Ambadra suriga (Borneo and
Wallacea) and Quadricalcarifera nigribasalis (Sumatra, Borneo, China,
Only 13 of the species are widespread in the Indo-Australian tropics, and a
similar number extend from the Himalaya to Sundaland. About 30 of the species
belong to morphologically isolated groups, many monobasic, restricted to
Sundaland or extending through Sundaland to Wallacea. The monobasic groups
include the genera Besida, Blakeia, Cerasana, Medanella, Pseudohoplitis,
Caschara and Rodneya, and the species Turnaca spinifera,
Neostauropus major and Ginshachia sumatrensis. Largely allopatric
arrays of species with a similar distribution occur in Hunyada, Pantanopsis,
Pseudostauropus, Stauroplitis, Sagamora and the Clostera dorsalis group.
Other groups with this geography include two or more sympatric species, namely Brykia,
Parasinga (though further species from outside the area may await
transference from Quadricalcarifera), the Clostera bramah group
and the Turnaca punctata group. Greatest complexity is shown by the Erconholda
group of species within Phalera where there is as much diversity in
the Philippines as in Sundaland.
The single association with New Guinea is shown by Euhampsonia gigantea, though
the genus Omichlis has its greatest diversity in New Guinea and the
Sundaland species may prove to have a sister relationship to the east; the male
genitalia are very different from those of the Himalayan species O.
There are two Oriental/Melanesian sister pairs of the sort described by Holloway
(1982b). One is found in the genus Archigargetta, and Blakeia
marmorata may have as its sister species Pseudo gargetta diversa Bethune
Baker (New Guinea, Solomons).
The Bornean species of Cerura, Oxoia, Teleclita and Allata, together
with those of the Phalera grotei group, belong to arrays of species,
largely allopatric, that extend throughout the Indo-Australian tropics, the
species falling into localised distribution categories as discussed by Holloway
(1982a), e.g. mainland Asian, Sundanian, Wallacean, Lesser Sundan, and
A number of endemic or Sundanian species have sister-relationships with mainland
Oriental species as follows:
Dudusa vethi: D. nobilis
Walker (S. China, Taiwan).
Gargetta costigeroides: G. costigera
Epistauropus apiculatus: E. vinaceus Moore
Tensha postobscura: T. striatella Matsumura
Saliocleta nannion: Saliocleta sp.
(India) + S. barasamphia Schaus (Philippines).
Liparopsis sundana +
L. dympna Schaus (Philippines): L. postalbida Hampson (N.E.
Fentonia sumatrana +
F. helena: F. ocypete Bremer (Himalaya to Amur) + F. excurvata Hampson
Fentonia bipunctus +
F. talboti: F. notodontina Rothschild (N.E,. Himalaya).
Notodontella nieuwenhuisi: N. ferrifusa Dudgeon
Chadisra borneensis: C. bipars Walker
or C. albobrunnea Rothschild (Himalaya).
Pseudoteleclita flavisticta: P. centristricta Hampson (Sri Lanka, India).
Ginshachia bronacha: G. gemmifera Moore
(N.E. Himalaya, China, Taiwan).
Clostera angularis: C. transecta Dudgeon
Plusiogramma aurosigna: P. argentata Oberthur
Many of these sister-patterns are reflected at a subspecific level by
species such as Formofentonia orbifer (cf. Ginshachia),
Quadricalcarifera nigribasalis (cf. Tensha), Porsica ingens (cf. Clostera)
and Tarsolepis rufobrunnea (cf. Pseudoteleclita).
Several other species are related to taxonomically unexplored mainland
Oriental complexes, such as Euhampsonia roepkei, Phalera javana, Phalera
acutoides, Antiphalera sumatrana, Peridea albipuncta, Disparia sundana and Ramesa
tripunctata. Other species belong to genera with a more even distribution of
diversity throughout the Oriental tropics, such as Gargetta, Hyperaeschrella and
Hexafrenum. The Chadisra complex extends throughout the
Indo-Australian tropics, and Porsica and Ambadra have their
centres of diversity in Sundaland. Quadricalcarifera is very diverse
throughout the Indo-Australian tropics, particularly in montane habitats.
Relationships within Saliocleta, Norraca and Micromelalopha await
Clostera fulgurita and C. restitura are widespread Oriental species that have
similarly widespread Palaearctic counterparts, and the latter also has a
relative in Melanesia.
There are three instances where a Bornean endemic has as a sister species a more
widely distributed yet sympatric species: Benbowia kiriakoffi/virescens;
Turnaca altipunctata/punctata; Clostera bramoides/bramah.
Comparisons with Sumatra
Of the 61 Bornean species restricted to the Sunda Shelf, 31 are shared
exclusively with Sumatra yet only two are shared exclusively with the Malay
Peninsula and none exclusively with Java. No doubt further collecting in the
last two localities will change this picture but the difference is sufficiently
striking for one to suggest that it will persist.
The list of Sumatran species by Bender & Dierl (1977) totals 154 species.
Thus, the recorded fauna would appear to be significantly larger than that of
Borneo despite the recording of several species twice through synonymy or errors
in past literature (e.g. the record of Antiphalera bilineata probably
refers to A. sumatrana, also recorded, though in combination with Grangulina).
Species shared with Sumatra comprise roughly 80% of the Bornean total,
suggesting close association between the two faunas in the past either directly
through exchange or indirectly through gaining the same species when mainland
Asian taxa were extending their ranges. It is interesting, therefore, to
consider the 25 or so Sumatran species not represented in Borneo that are listed
by Bender & Dierl, and also the relationships of the Bornean endemics.
The majority of these Sumatran species are found also in the Himalaya and a few
are known also from Java and Bali; a distribution typical of the 'Sumatran
track' pattern recognised by van Steenis for montane plant taxa (Holloway 1970
and in press a). Representation of this distribution type is very much
weaker in Borneo than Sumatra. It is not clear from the list of Bender &
Dierl which species are montane but greater penetration by montane mainland taxa
might be one explanation for the larger fauna. Species concerned are Baradesa
omissa Rothschild, Omichlis rufotincta Hampson (though the genus has
its centre of diversity in New Guinea), Hyperaeschra ochropis Hampson, Chloroceramis
viridinota Hampson, Metaschalis disrupta Moore and Rosama
In addition, Formotensha basalina Gaede has close relatives in the
Himalaya and elsewhere and Betashachia angustipennis Matsumura,
described from Taiwan, has a distinct subspecies in Sumatra, a remarkably
disjunct distribution that may represent the extreme of the pattern shown by Quadricalcarifera
nigribasalis and the genus Tensha. Perhaps further collecting in
Borneo and the Philippines will 'fill in the gaps' in this pattern and
reveal it as 'Luzon track' montane distribution (Holloway in press a). Mesophalera
obliterata Kiriakoff appears to be close to M. lundbladi Kiriakoff
from northern Burma.
Another group of Oriental species with similar extension to Sumatra, and
sometimes Java, falls within the Pydna Walker group of genera recognised
by Kiriakoff (1962), again much richer in Sumatra than Borneo. The species in
Sumatra not recorded from Borneo are Antheua servula Drury, a legume
feeder in a genus somewhat distinct from the rest, Poncetia albistriga Moore,
Norraca longipennis Moore (records of the Philippine Oraura ordgara Schaus
may refer to longipennis), Mimopydna essa Swinhoe, Niganda
strigifascia Moore, Pydnella rosacea Hampson, Pydnella galbana Swinhoe
and Hypambadra delineivena Swinhoe (see Holloway 1982 a: 208; the
monotypic genus Hypambadra Kiriakoff may be allied to Tensha Matsumura,
indicated by similar modification of the uncus, and the pair a sister-group to Ambadra
Moore). Host-plant records for the group are mainly of palms, grasses and
bamboos. Therefore the species may be characteristic of open or disturbed
habitats where such plants are well represented. Such habitats may have been
more extensive from the seasonally arid parts of Burma through Sumatra to Java
during dry periods of the Pleistocene glaciations, more so than in Borneo. Phalera
combusta Walker, also a grass feeder, falls into this category too.
The majority of species supposedly endemic to Sumatra listed by Bender &
Dierl have now been recorded from Borneo, or have Bornean sister species or
close relatives. Exceptions are Neopheosia albiplaga Gaede (the
relationship with N. fasciata needs to be reexamined), Corinella
vittata Gaede (affinities with Allodonta, Hexafrenum and allies), and
Closteroides dorsalis Kiriakoff, none of which have close relatives in
There are more endemic Quadricalcarifera species in Sumatra than Borneo.
The genus contains numerous species, mainly montane, from the Himalaya to New
Guinea, and is clearly in need of revision.
Most Bornean endemics are closely related either to species found in Sumatra or
to widespread Sundanian species. Exceptions are Phalera styx, with its
closest relatives in the Philippines, Turnaca spinifera, with
relationships obscure but possibly overlooked in Sumatra, Quadricalcarifera
trioculata, possibly related to more easterly species in Sulawesi or New
Guinea, Chadisra borneensis, with a Himalaya sister species but itself
possibly overlooked in Sumatra, and Micromelalopha cornutijuxta, possibly
the sister to the more widespread leucorhetha group.