::The Moths of Borneo::

Superfamily COSSOIDEA (48 species)

Family BRACHODIDAE (not covered)

Family COSSIDAE (37 species)

Subfamily COSSINAE (14 species)

Groenendaalia kinabaluensis Gaede (Part 1, p. 24, under Cossus). S. Burma, Thailand (Yakovlev, 2004a), Sundaland. Lowland to lower montane. Note 1. Note 1 . Groenendaalia Yakovlev. Yakovlev (2004a) erected this genus for kinabaluensis Gaede on the grounds of its unusual facies, combined with having simple male antennae, and, in the male genitalia, a robust process on the valve costal margin, a reduced saccus and a rather elongate, U-shaped juxta.

Aholcocerus verbeeki Roepke (Part 1, p. 24, under Cossus). Java, Bali, Borneo. (Lowland?). Note 2 Note 2 . Aholcocerus Yakovlev (type species ronkayorum Yakovlev, Pakistan). Yakovlev & Witt (2009) transferred verbeeki toAholcocerus. The male antennae are prismatic as in Holcocerus. The forewings have the basal zone tinged a creamy yellow. In the male genitalia, the uncus tends to be rounded, the valves have their apex relatively acute, and there is a large, rather corrugated triangular process extending over the centre of the valve costa. Yakovlev & Witt (2009) described A. ihleorum Yakovlev & Witt from Thailand and noted occurrence of the genus on the Nicobar Is..

Holcocerus chloratoides Holloway (Part 1, p. 24, under Cossus). Borneo, Peninsular Malaysia (B). (Lowland). Note 3 Holcocerus Staudinger. Schoorl (1990: 54‑55) treated this genus as distinct, but his analysis placed it as sister‑group to Cossus. The male and female antennae are prismatic in Holcocerus (referred to as filiform in Part 1) but each flagellomere is unilobed in Cossus. In this, both genera show a reduction, treated by Schoorl as synapomorphy, from the plesiomorphic bipectinate condition. There is also a synapomorphy in the basisternum of the stenopleural region of the mesothorax: this sclerite is shortened and rounded posteriorly. Schoorl placed two Bornean species in Holcocerus, and the antennae ofchloratoides Holloway indicate it may also belong in this genus, comb. n. H. chloratoides has been recorded from Peninsular Malaysia (FRIM colln). Holcocerus as recognised by Schoorl extends to Eurasia and Africa, but is not found further east than Sundaland in the Oriental region..

Paracossus chloratus Swinhoe (Part 1, p. 25, under Cossus). Sundaland, Wallacea. Lowland. Note 4 Paracossus Hampson. This genus, as circumscribed by Schoorl (1990), contains species where the male antennae are variously unilobed, bilobed, or more pectinate, and where the female antennae, where known, are prismatic. The only autapomorphy indicated in the cladistic analysis (Schoorl, 1990: 54‑55) is reduction in length of the fringes of the wing margins. The labial palps have scaling that is short ventrally and moderately long dorsally. The genus is distributed from India to Sulawesi (16 species listed by Schoorl), with one undescribed species in New Guinea. It appears to be sister‑group (Schoorl, 1990: 55) to four New World genera..

Paracossus speideli Holloway (Part 1, p. 26, under Cossus). Sundaland. (Lowland).

Paracossus microgenitalis Yakovlev (Plate 6, Fig 87). Endemic. (Montane forest). Note 5 Note 5 . Yakovlev (2004b: 374) described a new Bornean species, Paracossus microgenitalis Yakovlev (Plate 6, Fig 87). The holotype was from 1450m at Apin Apin on G. Trus Madi in Sabah. Two further specimens from Sabah have been seen, both males: 620m in undisturbed Agathis hill forest at Ginseng Camp, Maliau Basin (in FRC, Sepilok); 900m in stunted hill forest on Bukit Monkobo (in BMNH). The forewings are a satiny white, striated and shaded with dark greyish brown as illustrated. This shading is strongest on the dorsum with a strong extension into the medial zone that then extends more weakly and obliquely to the apex..

Roepkiella javanus Roepke (Part 1, p. 26, under Cossus). Sundaland. Lowland. Note 6 Note 6 . Roepkiella Yakovlev & Witt. Paracossus thaika Yakovlev was described by Yakovlev (2006) from S. Thailand. The illustration in the original description appears externally similar to chloratus Swinhoe and javanus Roepke, but no comparison was made with these species (Yakovlev, 2006), only with two others that are considerably larger. The male genitalia appear to be similar to those of chloratus.

Subsequently (Yakovlev & Witt, 2009), thaika and javanus have been placed in a new genus, Roepkiella Yakovlev & Witt, together with five other Oriental species, including two more described in Paracossus by Yakovlev (2006), and celebensis Roepke for Sulawesi. Yakovlev & Witt indicated that typical Paracossus males have bipectinate antennae, and certainly the taxa grouped in the genus by Schoorl show great variety of male antenna as mentioned in the earlier note. However, no mention is made of other taxa currently still in Paracossus, such as speideli and microgenitalis (see above), nor is chloratus referred to. One can only concur with the statement made by Yakovlev & Witt that ‘a complete revision of Indo-Malayan Cossinae must be undertaken’, but with a preference for ‘sooner’ rather than ‘later’, and employing cladistic methodology before any more genera are added to the plethora currently to hand..

Zeuzerocossus cinereus Roepke (Part 1, p. 26, under Cossus). Thailand, Sundaland. Lowland. Note 7 Zeuzerocossus Yakovlev. Yakovlev (2008b) placed cinereus in this new genus, defining it primarily on the unusual, partially bipectinate antennae as described in Part 1. The male genitalia and this antennal structure place the new genus close to Ronaldocossus Yakovlev (type species brechlini Yakovlev) from Sulawesi. On these grounds, Yakovlev placed both genera in a new tribe, the Zeuzerocossini, but did not indicate any features that were autapomorphic to the Cossini as a presumed sister-group; this action rendered the Cossini paraphyletic. Yakovlev recorded cinereus from central Thailand..

Isocossus cruciatus Holloway (Part 1, p. 28, under Cossus). Endemic. (Lowland). Note 8 Note 8. Isocossus Roepke. This genus also has bipectinate antennae in the male, but the rami only have hairs irregularly distributed on the proximal side; in Paracossus there are two rows of hairs on the inner margin and two rows on the outer one in species where the antennae are bipectinate (Schoorl, 1990: 97). However, this antennal character is shared with all genera in Section 3 of Schoorl (1990: 68‑69). The condition of the fringes of the wing margins and the scaling of the labial palps seem similar to that in Paracossus (cf. p. 54), and other characters appear ambiguous across the two cladograms and amongst the definitive characters listed for each Section. The only clear distinction would therefore appear to be in the male antennae. Two of the Bornean species of Isocossus described in Part 1 have subsequently proved to be conspecific following dissection of a long series of specimens taken in Peninsular Malaysia (Dr Tan Man Wah, pers. comm.). They are I. telisai and I. seria, the former having page priority, syn. n.

Schoorl (1990: 46) noted that Cossus rufipecten Holloway had males with completely bipectinate antennae, and was unsure whether this species should be assigned to Paracossus. The original description in Part 1 compared rufipecten with vandeldeni Roepke which it resembles most closely in size and facies. Schoorl placed vandeldeni in Isocossus Roepke, its genus of original description. Yakovlev (2006) formally assigned rufipecten to Isocossus.

This raises the question of the polarity of the two antennal conditions. It is notable that Schoorl (1990) referred the Bornean species cruciatus Holloway, retak Holloway and telisai Holloway to Paracossus, whereas Yakovlev (2006) transferred these to Isocossus. They are listed under the latter here, but the situation requires further analysis on a more stringent cladistic basis.

Yakovlev (2006) described I. stroehli Yakovlev from Sumatra and Peninsular Malaysia. This appears externally to resemblerufipecten closely (see particularly fig. 68e in Part 1), butrufipecten is not mentioned in the diagnosis, nor vandeldeni Roepke, though these are listed in a short review of species constituting the genus. The male genitalia appear to be similar to those of rufipecten, so it is likely that stroehli is conspecific..

Isocossus retak Holloway (Part 1, p. 28, under Cossus). Endemic. (Upper montane).

Isocossus telisai Holloway (Part 1, p. 28, under Cossus; now includes seria Holloway). Borneo, Peninsular Malaysia. (Lowland). Note 8 Note 8 . Isocossus Roepke. This genus also has bipectinate antennae in the male, but the rami only have hairs irregularly distributed on the proximal side; in Paracossus there are two rows of hairs on the inner margin and two rows on the outer one in species where the antennae are bipectinate (Schoorl, 1990: 97). However, this antennal character is shared with all genera in Section 3 of Schoorl (1990: 68‑69). The condition of the fringes of the wing margins and the scaling of the labial palps seem similar to that in Paracossus (cf. p. 54), and other characters appear ambiguous across the two cladograms and amongst the definitive characters listed for each Section. The only clear distinction would therefore appear to be in the male antennae. Two of the Bornean species of Isocossus described in Part 1 have subsequently proved to be conspecific following dissection of a long series of specimens taken in Peninsular Malaysia (Dr Tan Man Wah, pers. comm.). They are I. telisai and I. seria, the former having page priority, syn. n.

Schoorl (1990: 46) noted that Cossus rufipecten Holloway had males with completely bipectinate antennae, and was unsure whether this species should be assigned to Paracossus. The original description in Part 1 compared rufipecten with vandeldeni Roepke which it resembles most closely in size and facies. Schoorl placed vandeldeni in Isocossus Roepke, its genus of original description. Yakovlev (2006) formally assigned rufipecten to Isocossus.

This raises the question of the polarity of the two antennal conditions. It is notable that Schoorl (1990) referred the Bornean species cruciatus Holloway, retak Holloway and telisai Holloway to Paracossus, whereas Yakovlev (2006) transferred these to Isocossus. They are listed under the latter here, but the situation requires further analysis on a more stringent cladistic basis.

Yakovlev (2006) described I. stroehli Yakovlev from Sumatra and Peninsular Malaysia. This appears externally to resemblerufipecten closely (see particularly fig. 68e in Part 1), butrufipecten is not mentioned in the diagnosis, nor vandeldeni Roepke, though these are listed in a short review of species constituting the genus. The male genitalia appear to be similar to those of rufipecten, so it is likely that stroehli is conspecific..

Isocossus marginimaculatus sp. n. (p. 280, Plate 6, Fig 86). Endemic. (Logged lowland forest). Note 9 Note 9 .Isocossus marginimaculatus sp. n. (Plate 6, Fig 86). GG 15mm, 17mm. The male antennae are filiform. The forewing ground is a rather dirty white that is irrorated with a pale, dull, reddish brown extensively over the basal two‑thirds and more narrowly at the margin. The basal brown area is heavily mottled with dark grey that almost obscures it, but is much reduced in the submarginal zone of ground colour. The marginal brown zone contains a row of round black spots in the spaces at the very margin. These are present but fainter and greyer on the hindwing, which is otherwise completely irrorated with the reddish brown colour apart from sparse spotting of grey. The male gentalia are of the standard cossine type, perhaps most similar to those of I. telisai, but with the unusual wing facies being diagnostic. Holotype G. [SABAH]: Jln Tung‑hup, Telupid, 100m, in FRC, Sepilok. Paratype G. SABAH: Deramakot F.R., secondary forest (B), March 5, 1998, ( Ento Staff), in FRC, Sepilok. Geographical range . Borneo..

Isocossus rufipecten Holloway (Part 1, p. 27, under Cossus). Borneo, Sumatra. (Lowland and lower montane forest).

Isocossus vandeldeni Roepke (Part 1, p. 27, under Cossus). Sundaland. (Lowland).

Subfamily ZEUZERINAE (23 species)

Zeuzera indica Herrich-Schäffer (Part 1, p. 31). Indo-Australian tropics. Lowland.

Zeuzera caudata Joicey & Talbot (Part 1, p. 31). Indo-Australian tropics. Lowland. Note 10 Note 10. Yakovlev (2004a) and Yakovlev & Witt (2009) have treated Zeuzera rhabdota Jordan as a good species rather than as a subspecies of caudata, whereas Schoorl (1990) followed the latter course, also including celebensis Roepke as a synonym..

Zeuzera conferta Walker (Part 1, p. 32). Indo-Australian tropics. Lowland, especially mangrove.

Zeuzera coffeae Nietner (Part 1, p. 33). Indo-Australian tropics. Lowland, especially disturbed habitats.

Zeuzera lineata Gaede (Part 1, p. 34). Vietnam, Sundaland, Sulawesi. Montane.

Zeuzera borneana Roepke (Part 1, p. 34). Endemic. Lowland, (upper montane).

Zeuzera saikehi sp. n. (p. 281, Plate 6, Fig. 85). Endemic. Lower montane forest. Note 11 Note 11. Zeuzera saikehi sp. n. (Plate 6, Fig 85) GG 17mm. This is the smallest Bornean species and differs from the other smaller ones such as conferta Walker or coffeae Nietner in having the hindwing with a distinct tornal lobe somewhat as in caudata Joicey & Talbot, with the distal margin concave anterior to it. The wings have an unusual yellowish brown tinge over their basal halves. The blackish striae are relatively broad, somewhat as in lineata Gaede, but less numerous, and these are very faint on the hindwing, which is distinguished by a large blackish triangular mark on the anterior side of the tornal lobe. The genitalia have the uncus relatively long and slender as in caudata, but the valves are more ovate, less evenly rounded apically; the aedeagus is shorter.

HolotypeG. SABAH: Mile 10, Crocker Range, 1000m, 1983 (Saikeh) (in FRC, Sepilok).

Paratypes: 3GG as holotype with various dates (in FRC Sepilok, except one (cossid slide 421) in BMNH.

Geographic range:Borneo.

Xyleutes strix Linnaeus (Part 1, p. 35). Indo-Australian tropics. Lowland, especially alluvial, forest. Note 12 Note 12. Xyleutes Hübner. Schoorl (1990) restricted this genus to strix Linnaeus, persona le Guillou, and keyensis Strand (Kai, Tanimbar, ?New Guinea). It is defined on elevation of the frons, very short marginal fringes to the wings and the relative size of two sclerites at the base of the forewing (a large humeral plate). The two species in Borneo have an elongate black mark at two thirds from the base of the forewing, just at the apex of the cell. Schoorl noted a larval host record of Sesbania (Leguminosae) for strix.

Xyleutes persona le Guillou (Part 1, p. 35). Indo-Australian tropics to New Guinea. Lowland.

Chalcidica mineus Cramer (Part 1, p. 36, under Xyleutes). Indo-Australian tropics. Lowland. Note 13 Note 13. Xyleutes Hübner. Schoorl (1990) restricted this genus to strix Linnaeus, persona le Guillou, and keyensis Strand (Kai, Tanimbar, ?New Guinea). It is defined on elevation of the frons, very short marginal fringes to the wings and the relative size of two sclerites at the base of the forewing (a large humeral plate). The two species in Borneo have an elongate black mark at two thirds from the base of the forewing, just at the apex of the cell. Schoorl noted a larval host record of Sesbania (Leguminosae) for strix.

Bergaris malayica Roepke (Part 1, p. 38, under Xyleutes). Sundaland. Lowland forest. Note 14 Note 14. Bergaris Schoorl. This genus was described to include five species that range from Vietnam to New Guinea. No autapomorphies are indicated for this genus in the cladistic analysis (Schoorl, 1990: 141). The wing fringes are short and the humeral plate of the forewing is large.

Bergaris lutescens Roepke (Part 1, p. 39, under Xyleutes). Sundaland. Lowland.

Rapdalus pardicolor Moore (Part 1, p. 38, under Xyleutes). N.E. Himalaya to Sundaland. (Lowland alluvial forest). Note 15 Note 15. Rapdalus Schoorl. This is a monobasic genus, the type species pardicolor Moore being small with an unusually patterned forewing. Vein R1 is close to the forewing areole, and the labial palps are slender.

Hermophyllon anceps Snellen (Part 1, p. 39, under Xyleutes). Sundaland, Philippines. (Lowland alluvial forest). Note 16 Note 16. Hermophyllon Schoorl. Only the type species, anceps Snellen, is included, and Schoorl (1990) noted Xyleutes plesseni Schultze (Philippines) as a synonym. The genus was associated with Zeuzera in the phylogenetic analysis (Schoorl, 1990: 161), and also Tarsozeuzera Schoorl, all these genera having the labial palps 2‑segmented. The facies is very different from that of Zeuzera.

Tarsozeuzera fuscipars Hampson (new record, Plate 6). N. India, Thailand (Yakovlev, 2004a), Borneo, Peninsular Malaysia. (Lowland). Note 17 Note 17. Tarsozeuzera Schoorl. Schoorl (1990) defined this genus on thickening of the first tarsomere of the proleg and/or hindleg. The facies is distinctive, as illustrated for T. fuscipars Hampson in Plate 6. Barlow (1982) recorded T. fuscipars Hampson for Borneo, but this was not included in the Part 1 treatment because a voucher specimen for Borneo could not be located. This has now been remedied as a male from Tongod in Sabah was noted in FRC, Sepilok. The genus contains T. fuscipars, T. kochi Semper (the type species; Philippines, Sulawesi) and two undescribed African species. Barlow (1982) noted Gliricidia (Leguminosae) as a host plant for fuscipars.

Panau quarlesi Roepke (Part 1, p. 37, under Xyleutes). Endemic. (Lowland). Note 18 Note 18. Panau Schoorl. This genus is placed in a sister‑relationship to the next three in the phylogenetic analysis (Schoorl, 1990: 164‑175). The species are all of rather similar facies, most with a whitish marking on the forewing and with dark streaks distally between the veins. Schoorl found it otherwise difficult to define the genus. Possibly eleven species occur (Schoorl, 1990: 165‑166) through the Indo‑Australian tropics from Sundaland to New Guinea, with three in Borneo.

Panau stenoptera Roepke (Part 1, p. 37, under Xyleutes). Vietnam, Laos, Sundaland. Lowland.

Panau adusta Roepke (Part 1, p. 38, under Xyleutes). Vietnam, Sundaland. Lowland forest.

Duomitus ceramica Walker (Part 1, p. 36, under Xyleutes). Indo-Australian tropics. Lowland, especially alluvial, forest, also plantations. Note 19 Note 19. Duomitus Butler. Schoorl (1990) treated this genus as monobasic, including only the well-known teak pest, D. ceramica Walker. It shares with the next two genera tegulae that are tapering and upturned ventero‑posteriorly, and has a characteristic ligneous forewing facies. Chey (1996) has described its biology and pest status in Sabah.

Skeletophyllon euphyes West (Part 1, p. 39, under Xyleutes). Thailand (Yakovlev, 2004a), Sundaland, Wallacea. (Mangrove). Note 20 Note 20. Skeletophyllon Schoorl. This genus contains three medium‑sized species from Sundaland to New Guinea with facies as in the Bornean euphyes West and with slender labial palps. Vein R1 of the forewing is proximal to the areole. The Bornean species of this and the next genus appear to have some affinity for mangrove.

Trismelasmos dictyograpta Roepke (Part 1, p. 40, under Xyleutes). Endemic. (Mangrove, swamp and heath forest). Note 21 Note 21. Trismelasmos Schoorl. Schoorl (1990: 170) included 25 species in this new genus, most undescribed and the majority from the Australasian tropics, especially New Guinea where 18 species are listed, but with species as far east as the Solomons. There may be some association with mangrove. The two Sundanian species occur in Borneo and have forewing facies that may typify the genus: striae and spots of blackish brown on white, with blocks of the former colour around the margin. The third segment of the labial palps is short, ovate.

Skeletophyllon friedeli Yakovlev (2006) from S. Thailand is very similar in its unusual wing pattern to T. dictyograpta Roepke.

Trismelasmos maculatus Snellen (Part 1, p. 40, under Xyleutes). Sundaland, Lesser Sundas, Sulawesi. Lowland, especially mangrove, forest, (lower montane forest).

Relluna nurella Swinhoe (Part 1, p. 30, under Azygophleps). N.E. Himalaya, S.W. China, N. Burma, N. Vietnam, Thailand (Yakovlev, 2004a, 2008a), Sundaland, Palawan. (Lowland). Note 22 Note 22. Relluna Schoorl. This genus is monobasic, the type species being nurella Swinhoe. The three Indian species that Holloway (1982; Part 1) noted as having facies similar to that of Phragmataecia Newman, but with male genitalia that were closer to those of Azygophleps Hampson, may all belong to Phragmacossia Schawerda, though only parvipunctus Hampson was placed there with confidence by Schoorl (1990: 106). Yakovlev (2008a) expanded the known distribution of nurella and separated material from S. Burma and Sundaland as ssp. wallacei Yakovlev from typical material (N.E. Himalaya, N. Burma, S.W. China, N. Vietnam, N. Thailand). He recorded the species additionally from Palawan, possibly also ssp. wallacei.