The information on distribution and habitat preference is summarised in the following table and in the Checklist:

Distribution type

Total Ecological preference  
montane  lowland wide


18 6 9 3


61 (*31,**2) 12 36 13

Sundaland +Wallacea

12 - 8 4

Sundaland + New Guinea

1 - 1 -

Sundaland + N.E. Himalaya
(+ Wallacea) (+ Taiwan)

15 4 9 2

Oriental tropics to Sundaland

9 5 4 -

Oriental tropics to Wallacea

3 1 2 -

Oriental tropics to New Guinea

1 - 1 -


2 1 1 -


122 29 71 22

* Shared with Sumatra only.
** Shared with Peninsular Malaysia only.

The two species in the ‘Other’ category are Ambadra suriga (Borneo and Wallacea) and Quadricalcarifera nigribasalis (Sumatra, Borneo, China, Taiwan).

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. rufotincta Hampson.

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 Melanesian.

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
Walker (India).
Epistauropus apiculatus: E. vinaceus
Moore (India).
Tensha postobscura: T. striatella
Matsumura (Taiwan).
Saliocleta nannion: Saliocleta
sp. (India) + S. barasamphia Schaus (Philippines).
Liparopsis sundana
+ L. dympna Schaus (Philippines): L. postalbida Hampson (N.E. Himalaya, Taiwan).
Fentonia sumatrana
+ F. helena: F. ocypete Bremer (Himalaya to Amur) + F. excurvata Hampson (N.E. Himalaya).
Fentonia bipunctus
+ F. talboti: F. notodontina Rothschild (N.E,. Himalaya).
Notodontella nieuwenhuisi: N. ferrifusa
Dudgeon (N.E. Himalaya).
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 (N.E. Himalaya).
Plusiogramma aurosigna: P. argentata
Oberthur (China).

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 resolution.

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 plusioides Moore.

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 Borneo.

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.

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