taxonID	type	description	language	source
3027941DFFD3FFA7FEF2AFD5FB5DF8F1.taxon	description	(Figs. 1 A, C, E, F; 2 A – C, G – I).	en	Tang, Hongqu, Cranston,, Peter S. (2025): Paraskusella Cranston, 2018 and Kribiodosis Kieffer, 1921 (Diptera: Chironomidae: Chironomini): two new species from Borneo described, allowing incorporation into a multi-gene molecular phylogeny. Raffles Bulletin of Zoology 73: 34-42, DOI: 10.26107/RBZ-2025-0003
3027941DFFD3FFA7FEF2AFD5FB5DF8F1.taxon	materials_examined	Material examined. Holotype male, slide mounted in Euparal, MALAYSIA: Sarawak, Gunung Mulu National Park, Lupar tributary of Melinau River, 75 m a. s. l., 04 ° 1 ′ 48.8 ″ N 114 ° 49 ′ 4.4 ″ E, 12 June 2023, light trap, coll. H. Q. Tang (EJNU). Paratype male, as holotype (ANIC), teneral male in alcohol, as holotype except 11 July 2024 (SFC)	en	Tang, Hongqu, Cranston,, Peter S. (2025): Paraskusella Cranston, 2018 and Kribiodosis Kieffer, 1921 (Diptera: Chironomidae: Chironomini): two new species from Borneo described, allowing incorporation into a multi-gene molecular phylogeny. Raffles Bulletin of Zoology 73: 34-42, DOI: 10.26107/RBZ-2025-0003
3027941DFFD3FFA7FEF2AFD5FB5DF8F1.taxon	etymology	Etymology. From the type locality, Sarawak, the largest state of Malaysia on the island of Borneo. To be treated as a noun in apposition.	en	Tang, Hongqu, Cranston,, Peter S. (2025): Paraskusella Cranston, 2018 and Kribiodosis Kieffer, 1921 (Diptera: Chironomidae: Chironomini): two new species from Borneo described, allowing incorporation into a multi-gene molecular phylogeny. Raffles Bulletin of Zoology 73: 34-42, DOI: 10.26107/RBZ-2025-0003
3027941DFFD3FFA7FEF2AFD5FB5DF8F1.taxon	description	Description. Adult male (n = 2). Colour (Fig. 1 A). Generally yellow green, thorax without distinct vittae, postnotum brown. Foreleg with darker proximal band on femur, and stronger darkening on distal femur and all of tibia and tarsus. Middle and hind leg with distal darker band on femora, tibiae yellow, all tarsomeres browner apically. Total length 2.8 – 3.1 mm, wing length 1.35 – 1.45 mm. Head. Frontal tubercles absent. Flagellomeres 1 – 12, 370 – 390; flagellomere 13, 575 – 590; AR 1.51 – 1.55. Palpomeres 1 – 5: 25 – 30; 20 – 25; 87 – 100; 90 – 105; 140 – 148, Pm 3 and Pm 4 slightly thickened. Temporals 6 – 8, uniserial. Clypeus with 9 – 10 setae. Thorax with rounded scutum with weak median hump or smoothly rounded. Antepronotals 2 – 3; acrostichals 0; dorsocentrals comprising 4 anterior and 3 – 4 more posterior; prealars 2 – 3. Scutellum with 4 setae. Wing (Fig. 1 C) appears ‘ smoky’ with some darkening alongside veins, anal lobe weak. VR 1.17 – 1.25, Setation: R 0, R 1 0, R 4 + 5, 0, squama bare. Legs (Fig. 1 A, 2 A – C). Fore tibia with round scale occupying half circumference. Spurs on mid and hind combs straight, 25 – 40 long. LR 1 2.04 – 2.08, LR 2 0.60 – 0.62, LR 3 0.77 – 0.82; BV 1 1.39 – 1.41, BV 2 4.19 – 4.20, BV 3 2.88 – 2.89; SV 1 1.37 – 1.39, SV 2 3.65 – 3.78, SV 3 2.74 – 2.80. Hypopygium (Figs. 1 E, F; 2 G – I). Anal tergite bands separated medially to form incomplete V – shape, with two median anal tergite setae; 6 – 8 basal-lateral setae submarginal on tergite IX. Anal point (Fig 1 F, lateral view) spatulate, narrower in mid-section, 30 – 40 long from sclerotised base. Transverse sternapodeme broad, shallowly rounded anteriorly, with relatively strong oro-lateral projections. Phallapodeme indistinct. Gonocoxite c. 60 long, gonostylus c. 100 long. slender, inwardly curved, tapering to rounded point. Superior volsella apically crescent shaped, 35 – 40 long, without microtrichia, 2 medially-directed setae arising from distinct tubercle bases close to basal inner margin, with only 1 seta on outer margin. Median volsella (Fig. 2 H) c. 40 long, with strong stem, medio-distally with simple setae, medially broadened and apically recurved and tapered distally, mostly not extending beyond distal apex of superior volsella. Inferior volsella globular apically and extending beyond gonocoxite apex, microtrichiose apicoventrally, apically with medially and dorsomedially-directed, simple setae, without differentiated posteriorly directed strong seta. Female, pupa and larva unknown.	en	Tang, Hongqu, Cranston,, Peter S. (2025): Paraskusella Cranston, 2018 and Kribiodosis Kieffer, 1921 (Diptera: Chironomidae: Chironomini): two new species from Borneo described, allowing incorporation into a multi-gene molecular phylogeny. Raffles Bulletin of Zoology 73: 34-42, DOI: 10.26107/RBZ-2025-0003
3027941DFFD3FFA7FEF2AFD5FB5DF8F1.taxon	discussion	Remarks. Paraskusella sarawak, new species, conforms substantially to the morphological generic diagnosis for males of Paraskusella Cranston (2018). The thorax with only slight medio-dorsal bulge at most, is consistent with previously described species excepting Paraskusella hawkei Cranston, 2018 in which a protuberance is distinct (Cranston, 2018). The male of P. sarawak is separated from congeners by all the wing veins being bare.	en	Tang, Hongqu, Cranston,, Peter S. (2025): Paraskusella Cranston, 2018 and Kribiodosis Kieffer, 1921 (Diptera: Chironomidae: Chironomini): two new species from Borneo described, allowing incorporation into a multi-gene molecular phylogeny. Raffles Bulletin of Zoology 73: 34-42, DOI: 10.26107/RBZ-2025-0003
3027941DFFD3FFA2FC28AD14FA90FED1.taxon	description	(Figs. 1 B, D, G; 2 D – F, J – L)	en	Tang, Hongqu, Cranston,, Peter S. (2025): Paraskusella Cranston, 2018 and Kribiodosis Kieffer, 1921 (Diptera: Chironomidae: Chironomini): two new species from Borneo described, allowing incorporation into a multi-gene molecular phylogeny. Raffles Bulletin of Zoology 73: 34-42, DOI: 10.26107/RBZ-2025-0003
3027941DFFD3FFA2FC28AD14FA90FED1.taxon	materials_examined	Material examined. Holotype male, slide mounted in Euparal, MALAYSIA: Sarawak, Gunung Mulu National Park, Melinau Paku tributary stream, 75 m a. s. l., 04 ° 1 ″ 48.8 ″ N 114 ° 49 ′ 4.4 ″ E, 12 June 2023, light trap, coll. H. Q. Tang (EJNU). Paratypes 2 males, one on slide under 7 coverslips; hypopygium, wing and legs of partial additional DNA voucher specimen under 8 th coverslip; as holotype (ANIC); 2 males, as holotype, except 10 July 2024 (SFC).	en	Tang, Hongqu, Cranston,, Peter S. (2025): Paraskusella Cranston, 2018 and Kribiodosis Kieffer, 1921 (Diptera: Chironomidae: Chironomini): two new species from Borneo described, allowing incorporation into a multi-gene molecular phylogeny. Raffles Bulletin of Zoology 73: 34-42, DOI: 10.26107/RBZ-2025-0003
3027941DFFD3FFA2FC28AD14FA90FED1.taxon	etymology	Etymology. From the type locality, Gunung Mulu National Park. To be treated as a noun in apposition.	en	Tang, Hongqu, Cranston,, Peter S. (2025): Paraskusella Cranston, 2018 and Kribiodosis Kieffer, 1921 (Diptera: Chironomidae: Chironomini): two new species from Borneo described, allowing incorporation into a multi-gene molecular phylogeny. Raffles Bulletin of Zoology 73: 34-42, DOI: 10.26107/RBZ-2025-0003
3027941DFFD3FFA2FC28AD14FA90FED1.taxon	description	Description. Adult males (n = 2 – 3). Colour (Fig. 1 B) generally brown, thorax with darker brown vittae, scutellum yellow. All legs brown, with pale knee limited to articulated area. Abdomen with distinct dark brown posterior bands in segment I – VIII. Hypopygium brown in basal section and slightly pale in distal part. Total length 2.3 – 2.8 mm, wing length 1.1 – 1.3 mm. Head. Frontal tubercles absent. Flagellomere 1 – 12, 490 – 500 (n = 2); flagellomere 13, 370 – 380 m (n = 2). AR 0.74 – 0.78 (n = 2). Lengths (μm) of palpomeres (Pm) 1 – 5: 25 – 30; 30 – 35; 90 – 95; 80 – 90; 150 – 165, respectively (n = 2). Temporals 5 – 6, uniserial. Clypeus with 10 – 12 setae. Thorax with distinct scutal tubercle at 1 / 3 from antepronotum to scutellum. Antepronotals 0; acrostichals 0; dorsocentrals 4 – 6, sparsely distributed, with 1 – 2 humerals; tiny humeral pit present. Prealars 0. Scutellum with 2 setae. Wing (Fig. 1 D). Plain, wedge-shaped, anal lobe very reduced. VR 1.29 – 1.31. Setation: R 0, R 1 0, R 4 + 5 2 – 4, squama bare. Legs (Figs 1 B, 2 D – F). Length of fore-leg 4.0 – 4.3 mm, c. 7 × thoracic height. Fore tibia scale pointed, 30 - 35 long, subapex of tarsomere IV of fore-leg with 3 aligned stiff setae, 30 – 35 long. Long spurs of mid and hind combs 55 – 63 long, curved. LR 1 2.50 – 2.58, LR 2 0.95 – 0.98, LR 3 1.22 – 1.24; BV 1 2.10 – 2.24, BV 2 2.90 – 2.96, BV 3 2.46 – 2.48; SV 1 1.15 – 1.17, SV 2 2.44 – 2.52, SV 3 1.82 – 1.85. Hypopygium (Fig. 1 G, 2 J – L). Anal tergite bands separated medially, with 0 (? 1) median seta, 4 – 6 baso-lateral setae submarginal on tergite IX. Anal point narrow (4 wide at base) parallel-sided to slightly tapering to towards apex, c. 25 long from sclerotised base. Superior volsella (Fig. 2 K) curved medially, 25 – 30 long, with 3 medially-directed setae arising from basal-mid inner margin. Inferior volsella cylindrical, 70 long (inner margin) extending to mid-section of gonostylus, microtrichiose apicoventrally, apically with medially and dorsomedially-directed, simple setae, with 4 – 6 differentiated posteriorly directed strong seta. Gonostylus c. 50 long, slender, tapering to blunt point. Transverse sternapodeme broadly plate-shaped, bluntly rounded anteriorly, without oral projections. Phallapodeme simple, distinct. Female, pupa and larva unknown.	en	Tang, Hongqu, Cranston,, Peter S. (2025): Paraskusella Cranston, 2018 and Kribiodosis Kieffer, 1921 (Diptera: Chironomidae: Chironomini): two new species from Borneo described, allowing incorporation into a multi-gene molecular phylogeny. Raffles Bulletin of Zoology 73: 34-42, DOI: 10.26107/RBZ-2025-0003
3027941DFFD3FFA2FC28AD14FA90FED1.taxon	discussion	Remarks. K. mulu conforms substantially to the male generic diagnosis for Kribiodosis (Han et al., 2021) but is distinguished from congeners by the legs showing a limited pale ‘ knee’, and wing veins R and R 1 totally lacking setae (R 4 + 5 has few setae). The three stiff setae (‘ pseudospurs’) on Ta 4 in the fore-legs (Han et al., 2021: fig. 4 b), are unusual, possibly assisting in copulation. Similar structures in Chironomus claggi Tokunaga, 1964 are located on the mid – and hind leg tarsomeres (Yamamoto & Yamamoto, 2018), rather than on the fore leg as in Kribiodosis.	en	Tang, Hongqu, Cranston,, Peter S. (2025): Paraskusella Cranston, 2018 and Kribiodosis Kieffer, 1921 (Diptera: Chironomidae: Chironomini): two new species from Borneo described, allowing incorporation into a multi-gene molecular phylogeny. Raffles Bulletin of Zoology 73: 34-42, DOI: 10.26107/RBZ-2025-0003
3027941DFFD6FFA1FC55ABF4FC35F9D1.taxon	description	(Figs. 1 H – K, 3 A, C, E – I)	en	Tang, Hongqu, Cranston,, Peter S. (2025): Paraskusella Cranston, 2018 and Kribiodosis Kieffer, 1921 (Diptera: Chironomidae: Chironomini): two new species from Borneo described, allowing incorporation into a multi-gene molecular phylogeny. Raffles Bulletin of Zoology 73: 34-42, DOI: 10.26107/RBZ-2025-0003
3027941DFFD6FFA1FC55ABF4FC35F9D1.taxon	description	Labrum (Figs. 3 F) with SI on conjoined bases, seemingly shaped as smooth-edged fan, with minor setal fringe, and SII placed on a distinct pedestal, blade-like, apical ¼ with inner fringe, 30 – 35 long; SIII simple, 30 long; SIVa, b weak. 6 – 7 plumose chaetae. Seta premandibularis simple. Labral lamellae lie largely beneath SI, fringed. Pecten eipharyngis comprising three separated plates, each with 5 – 8 teeth; 7 – 8 apically finely plumose chaetulae laterales, 2 apically branched chaetulae basales. Premandible (Fig. 3 G) 70 – 80 long, with 4 sharply pointed teeth and modest brush. Mandible (Figs. 1 I, J, 3 E) 100 – 110, with dorsal tooth, pointed apical tooth and 3 pointed inner teeth. Pecten mandibularis comprising a 25 – 32 spine-like lamella, protruding to mandibular margin, other branches few, short, simple. Seta subdentalis (Fig. 3 E, inset) 30 – 35 long, arising from ventral surface, sickle-shaped, distally with comb, extending to notch delimiting apical mandibular tooth. Mola and inner margin smooth. Seta interna 4 – 5 branched, densely plumose, but short. Mentum (Figs. 1 H, 3 A) 60 – 75 wide, with 16 continuous pale gold teeth without a delimited ventromental component; central 2 teeth recessed, the 2 nd and 3 rd elevated thereafter evenly declining. Ventromental plate 80 – 85 wide, 40 – 45 deep, fan-shaped, with smoothly curved anterior margin, medially tapered to medially-directed (not upcurved) point, inter-plate distance subequal to two recessed median teeth; striae very fine, seemingly restricted to median, lateral / posterior plate, lappets / hooklets indistinct. Setae submenti simple. Abdomen (Fig. 3 I) without lateral or ventral tubules nor dorsal hump. Anterior parapod claws pale, dense, fine, simple; posterior parapods claws golden, simple. Procercus pale yellow, 2 × as high as wide (45 – 55 × 25 – 30), bearing 5 anal setae 400 – 750 long. Anal tubules elongate-ovoid (150 – 200 × 50 – 80), dorsal slightly shorter than ventral, subequal or slightly shorter than posterior parapods.	en	Tang, Hongqu, Cranston,, Peter S. (2025): Paraskusella Cranston, 2018 and Kribiodosis Kieffer, 1921 (Diptera: Chironomidae: Chironomini): two new species from Borneo described, allowing incorporation into a multi-gene molecular phylogeny. Raffles Bulletin of Zoology 73: 34-42, DOI: 10.26107/RBZ-2025-0003
3027941DFFD6FFA1FC55ABF4FC35F9D1.taxon	discussion	Comments. The larva of Kribiodosis does not key beyond the cluster with 6 - segmented antenna and alternating distinctive Lauterborn organs on the apices of antennal segments 2 and 3 in Epler et al., 2013. The ‘ wedge-shaped’ (triangular) ventromental plate directs to couplet Zavreliella Kieffer, 1920 vs Lauterborniella Thienemann & Bause, 1913, both of which have case-bearing larvae and considered as sistertaxa on morphological grounds by Andersen et al. (2017). Of these, Kribiodosis resembles Zavreliella in having simple setae submenti (vs. plumose) and the dorsal head including a frontoclypeus (vs frons and isolated clypeus). A broad seta subdentalis with comb-like distal margin (Fig. 1 J) occurs in Kribiodosis and Lauterborniella, and in a different form in the distant Goeldichironomus Fittkau, 1965, Kiefferulus Goetghebuer, 1922 and Axarus Roback, 1980. Kribiodosis will not key to any Holarctic taxon even if restricted to those in which a ventromentum is not delimited (Apedilum). Southern hemisphere genera belonging to a ‘ Microtendipes – group’ show permutations of larval features. Included are Oukuriella Epler, 1986, Claudiotendipes Andersen, Mendes & Pinho, 2017, Conochironomus Freeman, 1961, Skusella Freeman, 1961, Paraskusella Cranston, 2018, Paraborniella Freeman, 1961 and Paucispinigera Freeman, 1959 (Cranston, 2020). In the Neotropics, Kribiodosis keys unequivocally to Zavreliella / Lauterborniella based on the sub-triangular ventromental plates in near contact medially (Silva et al., 2018). Claudiotendipes is excluded on these features. In the guide to Australian and New Zealand larvae (Cranston, 2019) despite the increased diversity, a similar conclusion is reached for identical reasons. Since the larva of Kribiodosis is unlikely to be case-bearing, judging by morphology, it can be excluded from Zavreliella or Lauterborniella. Larval keys indicate resemblance to several other taxa, amongst which is Beardius Reiss & Sublette, 1985, a diverse New World genus that includes species with the diagnostic antenna of the ‘ Microtendipes – group’. In both Zavreliella and Lauterborniella the basal antennal segment has a distinct ‘ antennal’ seta located subapically but this seta is lacking in Kribiodosis. In the plate of line drawings of Australian larval ‘ Zavreliella ’ (Cranston, 1996), the mentum and antenna of taxon ‘ K 1 ’ (K = Kakadu, northern Australia) and S 1 (S = Sydney) (Fig. 3 B, D), closely resemble those of Kribiodosis described here, in contrast to Australian Zavreliella marmorata (Wulp, 1859) on the previous page that conforms to the (‘ true’) Zavreliella larva (and pupa). Access to these specimens is currently unavailable due to databasing, reorganisation and relocation of slide collections at ANIC. These could confirm differences between genera as reported above. It is likely that Kribiodosis is to be found in tropical Australian waters. Unfortunately, as the associated pupae of both Kribiodosis species are unavailable to us, the presumed pupal type described in Cao & Tang (2017) is an unreliable individual association. Molecular results and discussion. Our molecular multi-gene analysis uses a set derived from exemplars of the ‘ Microtendipes – group’ of genera for which DNA data are available, including Conochironomus and Skusella, presumed relatives of Paraskusella. This is anchored with Riethia and Tanytarsus as outgroups which consistently have formed the sister group to tribe Chironomini (e. g., Cranston et al., 2011; Han et al., 2021). New sequences for Paraskusella and a second species of Kribiodosis from this report have been incorporated into an ongoing expanded study. Most genera represented by multiple species have been pruned to a single terminal, recognisable in Fig. 4 by the published codes that follow the genus name (e. g., Microtendipes TH 02, HW 013). Bayesian inference analysis revealed several strongly supported nodes identified by PP (posterior probability) = 0.99 – 1, indicated on Fig. 4 by ***, fewer weaker supported nodes (** PP = 0.95 – 0.98, * PP = 0.90 - 0.95) and with critical nodes lacking Bayesian support (PP <0.90 (unlabelled). Bootstrap values (72 – 100) are indicated only for nodes with Bayesian support> 0.90.	en	Tang, Hongqu, Cranston,, Peter S. (2025): Paraskusella Cranston, 2018 and Kribiodosis Kieffer, 1921 (Diptera: Chironomidae: Chironomini): two new species from Borneo described, allowing incorporation into a multi-gene molecular phylogeny. Raffles Bulletin of Zoology 73: 34-42, DOI: 10.26107/RBZ-2025-0003
3027941DFFD6FFA1FC55ABF4FC35F9D1.taxon	description	Biogeography. The revelation of ever-increasing taxa in the subfamily Chironominae, notably in the tribe Chironomini, showing African – (Australian) – Oriental distributions has been reported in Han et al. (2021) with regard to Kribiodosis. To the examples discussed there, we add Paraskusella, previously known from Africa and Australia, now reported from Borneo in the Oriental region. The closest relatives Conochironomus and Skusella each show the same distribution pattern, referred to as the tropical Gondwana track by Matile (1990) and discussed by Cranston (2005). Whether this remains a valid descriptor given recent discoveries of these taxa in (non-Gondwanan) oriental China is unclear at present. A robust and dated phylogeny that includes these taxa at species-level will be necessary to further explore this pattern.	en	Tang, Hongqu, Cranston,, Peter S. (2025): Paraskusella Cranston, 2018 and Kribiodosis Kieffer, 1921 (Diptera: Chironomidae: Chironomini): two new species from Borneo described, allowing incorporation into a multi-gene molecular phylogeny. Raffles Bulletin of Zoology 73: 34-42, DOI: 10.26107/RBZ-2025-0003
