taxonID	type	description	language	source
6D0F87C50114FFD4FF3FFDB9FEE3FCFF.taxon	materials_examined	Type species: Chironomus subvittatus Skuse, 1889: 246, by original designation. Other included species: Skusella pallidipes (Kieffer, 1921), Africa; Skusella freemani Harrison, 2002, Africa; Skusella silingae Tang sp. n., Asia.	en	Cranston, Peter S., Tang, Hongqu (2018): Skusella Freeman (Diptera: Chironomidae): new species, immature stages from Africa, Asia and Australia, and expanded distributions. Zootaxa 4450 (1): 41-65, DOI: 10.11646/zootaxa.4450.1.3
6D0F87C50113FFDEFF3FFB16FC25F89B.taxon	materials_examined	Material examined. Holotype. ♂ slide mounted in Euparal by Cranston: ‘ Walcha’ [AUSTRALIA, New South Wales], ‘ Ch. subvittatus ♂ type’, F. A. A. Skuse, ANIC [no collection date]. Other material (collected Cranston, deposited ANIC, unless otherwise stated). AUSTRALIA: Northern Territory, 5 Pe, nr. Narbaluk, Cooper Ck, 12 ° 18 ' S 133 ° 20 ' E, 27. v. 1988; 1 L, U. Magela Ck, Stoned Billabong, 12 ° 38 ' S 132 ° 53 ' E, 14. iv. 1989; 1 P ♂, U. Magela Ck, below Bowerbird Billabong, 12 ° 47 ' S 133 ° 03 ' E, 28. v. 1988; 2 L, Upper Magela Ck, above falls, 12 ° 47 ' S 133 ° 36 E, 15. iv. 1989; 1 L, 2 Pe. 1 Le / Pe / ♀, Wildman R. at Arnhem Highway Crossing, 12 ° 50 ' S 132 ° 01 ' E, 2. vi. 1988; 2 Pe, Hickey Ck, 12 ° 55 ' S 132 ° 50 ' S, 29. v. 1988; 2 Pe, S. Alligator R. at Kakadu Highway ford, 13 ° 17 ' S 132 ° 19 ' E, 26. v. 1988; 1 Pe, Jim Jim Ck, above falls, 13 ° 17 ' S 132 ° 51 ' E, 29. v. 1988 (Dostine) (ARRRI); 1 Pe, Jim Jim Gorge, 21. xi. 1989 (Dostine) (ARRRI); 1 Pe, Graveside Gorge, 13 ° 18 ' S 132 ° 32 ' E, 17. vii. 1988 (Dostine) (ARRRI # 165); 1 Pe, Koolpin Gorge, 13 ° 30 ’ S 132 ° 35 ’ E, 15 – 16. v. 1992; 1 ♂, Kambolgie Ck, 13 ° 32 ' S 132 ° 23 ' E, 25. v. 1988; 1 Pe, S. Alligator R. at Fisher Ck., 13 ° 33 ' S 132 ° 33 ' E, 24. v. 1988; 1 Le / P, 1 Pe, Gimbat, 13 ° 34 ' S 132 ° 35 ' E, 24. v. 1988; 2 ♂, Coronation Hill, 13 ° 35 ' S 132 ° 36 ' E, 4 / 5. vi. 1988; 3 Pe, [Nitmiluk N. P.] Katherine R., below gorge 1, 14 ° 20 ' S 132 ° 25 ' E, 21 – 22. v. 1992; 1 ♂, Caranbirini W. H., 33 km s. w. of Borroloola, 16 ° 19 ' S 136 ° 05 ' E, 3. xi. 1975 (Upton, Macarthur R. survey), 1 ♂, 36 km s. w. of Borroloola, 4. xi. 1975; 14 km s. w. Cape Crawford, 16 ° 47 ' S 135 ° 45 ' E, 25. x. 1975. Queensland, 1 ♂, 18 km w. Mt. Carbine, McLeod R., 16 ° 49 ' S 145 ° 00 ' E, 7. i. 1982 (Baehr) (ZSM); 1 ♂, 2 km e. Einasleigh, Einasleigh R., 18 ° 34 ' S 144 ° 06 ' E, 11 – 12. vi. 1993 (Baehr) (ZSM); 1 P / ♂, Lawn Hill N. P., Musselbrook Ck., upper pool, 18 ° 37 ' S 138 ° 08 ' E, 8. v. 1995; 1 ♂, Louie Ck., Lawn Hill Crossing, 18 ° 45.39 ' S 138 ° 31.15 ’ E, 17. v. 1995, swept; 1 ♂, same except light trap. New South Wales, 1 L, nr. Wooli, Lake Hiawatha, 2 m. benthic, 29 ° 48 ' S 153 ° 09 ’ E, 1. vii. 1971 (Timms); 1 L, Barrington Tops, Williams River at Rocky Crossing, 32 ° 07 ' S 151 ° 29 ' E, 8. xii. 1993 (Sydney Water); 1 L, Bardon, Dusodie, ' New Park', Chichester R., 32 ° 26 ' S 151 ° 69 ’ E, 8. xii. 1993 (Sydney Water); 4 Pe, nr Araluen, Deua N. P., Deua R., 35 ° 45 ' S 149 ° 57 ’ E, 29. iii. 1989. Victoria, 2 Pe, Little Beetle Lake, 37 ° 47 ' S 148 ° 25 ’ E, 17. xii. 1996 (Wright). Western Australia, 1 P (♂), Pilbara, Fortescue R., Fortescue Falls, 22 ° 29.00 ' S 118 ° 33.06 ’ E, CLM 26 M (Smith); Hamersley Range N. P., Circular Pool, Fortescue Falls, 22 ° 28 ' S 118 ° 33 ’ E, 23 – 24. iv. 1992; 1 Pe, Hamersley Gorge, 22 ° 29.00 ' S 118 ° 33.06 ’ E, 22. iv. 1992; 4 Pe, Millstream Chichester N. P., Chinderwarrinder Pool, 21 ° 35 ' 24 " S 117 ° 04 ’ 02 " E, 25. iv. 1992; 1 Pe, Crossing Pool, 21 ° 34 ' 22 " S 117 ° 05 ’ 02 " E, 24. iv. 1992; 1 Pe, 7 ♂, Millstream Chichester N. P., Fortescue R., below Homestead, 21 ° 33 ' S 117 ° 03 ’ E, 24 – 25. iv. 1992; 1 Pe, Willare Bridge, Fitzroy R., 17 ° 44 ' 08 " S 123 ° 38 ' 45 " E, 28. iv. 1992; 1 ♂, 30 km s. Warmun, Frog Hollow Ck, 17 ° 16 ' S 128 ° 03 ' E, 14 – 15.1984 (Baehr) (ZSM); Pe, Manning Gorge, 16 ° 39 ' S 125 ° 56 ' E, 30. v. 1992; 10 Pe, Drysdale R. crossing, 15 ° 41 ' S 126 ° 47 ' E, 1 – 2. v. 1992; 2 Pe, King Edward R., 14 ° 53 ' S 126 ° 12 ' E, 5 – 6. v. 1992; 1 ♂, 14 km s. e Kalumburu, CALM site # 4 / 3, 14 ° 25 ' S 126 ° 40 ' E, 3 – 6.1988 (Weir); 2 Pe, Old Boab Camp, Crystal Ck., 14 ° 29.42 ' S 125 ° 47.26 ' E, 3. v. 1992; 5 Pe, Mertens Falls, 13 ° 50 ' S 125 ° 43 ' E, 4 – 5. v. 1992. Non-Australian. BRUNEI, 2 Pe, Kuala Belalong, Kuala Belalong Field Study Centre, 04 ° 33 ' N 115 ° 09 ' E, -. viii. 1995.	en	Cranston, Peter S., Tang, Hongqu (2018): Skusella Freeman (Diptera: Chironomidae): new species, immature stages from Africa, Asia and Australia, and expanded distributions. Zootaxa 4450 (1): 41-65, DOI: 10.11646/zootaxa.4450.1.3
6D0F87C50113FFDEFF3FFB16FC25F89B.taxon	description	NEW CALEDONIA: 2 Pe, S. Prov., Chutes de Madeleine, 22 ° 14 ’ S 166 ° 52 ’ E, 1. ii. 2003; 2 L, 1 Pe, Mt Mou, Road Crossing, c. 200 m. asl, 22 ° 04 ’ S 166 ° 20 ’ E, 26. i. 2003. SINGAPORE: 2 L, Nee Soon swamp, site # 2, 01 ° 23 ' N 103 ° 48 ' 40 ” E, iii. 2009 (‘ NUS team’) (TMSI). Description. Adult male (n = 10, less for some legs). B. l. 2.9 – 3.9 mm, W. l. 1.4 – 1.8 mm. Fl 1 – 12 396 – 428. Fl 13 573 – 682, A. R. 1.40 – 1.59. Temporals 9 – 12 uniserial, some long; Clyp 14 – 26. Palp 2 – 5: 36 – 50, 136 – 208, 118 – 185, 159 – 318. Thorax with double hump, anterior shallower than posterior. L. aps absent. Dc 8 – 14, Pa 2 – 3, Scts 3 – 9. Wing. Plain. V. R. 1.01 – 1.12. Vein setation: R 11 – 22, R 1 9 – 17, R 4 + 5 13 – 22, squamals 0 – 5. Legs (Fig. 1 E). Apically curved spur on each mid and hind outer comb, inner comb with small, blunt spur barely extending beyond longest comb teeth. LR 1 2.0 – 2.2, LR 2 0.60 – 0.65, LR 3 0.83 – 0.88. BV 1 1.7 – 1.8, BV 2 3.2 – 3.5, BV 3 2.5 – 2.7. SV 1 1.25 – 1.3, SV 2 3.3 – 3.6, SV 3 2.46 – 2.5. Hypopygium (Figs 1 F, 1 H). TIX bare; anal point parallel-sided, with bluntly tapered apex. SVo (Fig. 1 H) ca. 50 – 60 µm, with 2 inner setae, 15 – 22 long, bases 5 – 12 µm apart, each arising from tubercle; 3 rd very fine seta, 10 long, present in some specimens either between, or distal to strong inner setae. Mvo stem ca. 30 µm, with 20 – 30 split semi-taeniate and simple setae. Gcx 113 – 160, Gst 96 – 108. Adult female (n = 1) as male, except: B. l. 2.9 mm, W. l. 1.5 mm. Antennal ratio 0.41; apical flagellomere 110. Clyp 23. Palp 2 – 5: 40, 190, 165, 300. Thorax with double scutal hump, anterior shallower than posterior. L. aps absent. Dc 11 – 12, Pa 3; Scts 11 – 12. Wing. V. R, 1.08. R 24, R 1 22, R 4 + 5 27; 1 squamal. Legs. LR 1 2.0, LR 2 0.57, LR 3 0.82. BV 1 1.7, BV 2 3.4, BV 3 2.6. SV 1 1 1.3, SV 2 3.5, SV 3 2.7. Genitalia as in Fig. 3. Pupa (n = 10). Length 4.0 – 4.8 mm. Exuviae pale, with only slightly darkened apophyses. Cephalothorax. Cephalic area smooth without tubercles, pale / hyaline frontal setae 30 – 40 µm. Thorax with weak scutal hump at anterior 1 / 3 with small scales mid-thorax alongside ecdysial line. Base of thoracic horn elongate oval, tracheal bundle densely packed, 20 x 15 µm (Fig. 4 C). Prealar tubercle rounded-triangular, darkened (Fig, 4 D). Abdomen (Fig. 5 A). Tergites II – VI with spinule patches laterally not reaching apophyses, and not extending to pleurae, narrower anteriorly than posteriorly; on VII and VIII spinule patches quite dense in anterior broad transverse band of spinules. Tergite II with row of 15 – 20 hooks extending 20 – 22 % of tergite width. Pedes spurii B moderately developed. Pleural fringe on segments III – VIII. Posterolateral ‘ comb’ on VIII with 10 – 14 teeth of relatively uniform size (Fig. 4 H). Larva (n = 2, reared). Head pale yellow with slightly darker golden-brown occipital margin, mandibles and mentum. Body length not measured. Head capsule length 450 µm, postmentum length 150 µm. Eyespots not visible. Dorsal surface of head as in Fig. 7 A. Antenna (Fig. 7 B) with segment lengths 87 – 90: 14 – 15: 38 – 42: 24 – 25: 5 – 63; AR 1.1. Lauterborn organs alternate on segment 3, 11 – 12 long; blade 50 – 62 long, extending to apex of segment 3. Labroepipharyngeal region as in Fig. 7 D, premandible (Fig. 7 E) 75 long, with 3 apical teeth. Mandible 110 – 126 long, as in Fig. 7 F. Mentum (Fig. 7 G) 82 – 90 wide; ventromental plates 108 – 112 wide, separated medially by 42 – 45. Abdomen. Anterior parapod claws densely clustered and very pale. Posterior parapod claws golden, squat, gently curved. Procercus 20 high by 20 wide, bearing 7 – 8 long pale anal setae.	en	Cranston, Peter S., Tang, Hongqu (2018): Skusella Freeman (Diptera: Chironomidae): new species, immature stages from Africa, Asia and Australia, and expanded distributions. Zootaxa 4450 (1): 41-65, DOI: 10.11646/zootaxa.4450.1.3
6D0F87C5011FFFDCFF3FFF0CFC25FE36.taxon	materials_examined	Material examined. 1 ♂, SUDAN [SOUTH SUDAN], Wupatong, nr Wau, 19. iii. 1955 (Reid) BM 1955 - 382 (NHM); 1 ♂ nr. Wau (Schorch) (NHM), 1 ♂, BELGIAN CONGO, Elisabethville [DR CONGO, Lubumbashi], iii. 1939 (Brédo) (NHM); 1 ♂, 1 ♀, IVORY COAST, no further data (Dejoux) (ZSM); 1 ♀, NIGERIA, Anambra State, Opi Lake 1 (= Ogelube Lake), 6 ° 45 ' 0 " N 7 ° 29 ' 30 " E, 4 – 5. xi. 1978, emergence trap (Hare) (NHM); 1 Pe (putative identity), same except 28. i. – 7. ii. 1978. Material mapped, not examined. Sudan [South Sudan]: Melut, Adok, Shambe; Nigeria: Katsina, Kankiya; Sierra Leone: Pepel; Gold Coast [Ghana]: Nangodi; French Cameroons [Cameroon]: Kribi; Belgian Congo [DR Congo]: Musosa; S. Rhodesia [Zimbabwe]: Salisbury [Harare].	en	Cranston, Peter S., Tang, Hongqu (2018): Skusella Freeman (Diptera: Chironomidae): new species, immature stages from Africa, Asia and Australia, and expanded distributions. Zootaxa 4450 (1): 41-65, DOI: 10.11646/zootaxa.4450.1.3
6D0F87C5011FFFDCFF3FFF0CFC25FE36.taxon	description	Description. Adult male (n = 5). B. l. 6.0 – 6.8 mm, W. l. 2.4 – 2.7. Antenna: Fl 1 – 12 405 – 445, Fl 13 1115 – 1240, A. R. 2.75 – 2.92. 19 – 14 temporals, 17 – 23 clypeals. Palps (2 – 5): 65 – 75, 145 – 180, 190 – 210, 300 – 370. Thoracic contour double humped; L. aps 4 – 5; ac 0, dc 9 – 13, pa 2 – 3, Scts 6 – 8. Vein setal counts R 7 – 11, R 1 16 – 17, R 4 + 5 17 – 22, VR 1.08 – 1.14. Wing patterned as in Fig. 1 B (female). Vein setal counts R 16, R 1 16 – 17, R 4 + 5 11 – 16. V. R. 1.18. V. R. 1.07 – 1.14. Squama bare. Legs: Fore-tibial scale large without spur, mid and hind tibial combs nearly fused, bearing 1 hooked spur. LR 1 1.65 – 1.80, LR 2 0.39 – 0.44, LR 3 0.68 – 0.74; BV 1 1.50 – 1.64, BV 2 3.30 – 3.50; BV 3 2.65 – 2.80. SV 1 1.41 – 1.54, SV 2 4.60 – 4.90, SV 3 2.88 – 3.00. Hypopygium (Fig 1 G, 1 I – J). Bands of tergite IX very faint, nearly meeting, lacking median setae, lateral to anal point TIX has several fine long seta. Superior volsella (Fig. 1 I) tapering to dorsally curved apex, with 2 long inner setae arising from large basal tubercles. Median volsella 50 µm, thin stemmed, with simple setae and taeniae (Fig. IJ). Gcx 240 – 300, Gst 165 – 202; H. R. 3.05 – 3.86. Adult female (n = 3, all under- or over-cleared; 1 slide label states ‘ dried out’). Body length c. 3.5 – 3.6 mm. Wing length 1.8 – 1.9 mm. Head 9 – 10 temporals, 14 – 15 clypeals. Antenna 110 – 130, 65 – 70, 70, 60, 150, AR 0.41 – 0.42. Palp lengths (2 – 5): 40, 210 – 220, 220, 310 – 315. Thorax. With double scutal hump, anterior larger than longer, shallower posterior. Setation: 1 L. aps, 9 – 10 dc, 3 pa, 3 – 5 scts. Wing (Fig. 1 B). Pattern similar to but fainter than male and figure (perhaps all “ bleached): with two broad separate darker bands each extending full width of wing. Vein setal counts R 23 – 28, R 1 21 – 28, R 4 + 5 33 – 46. V. R. 1.18. All legs damaged, all tarsomeres absent: pale, at most faintly darker at femoral-tibial junction. Genitalia. Similar to S. subvittata except coxosternapodeme IX thicker, darker and more sinuous; seminal ducts more sinuous / recurved, ending very close, perhaps fused in single opening. Seminal vesicles elongate-ovoid 105 – 120 long, maximum width 50 – 60, with strong neck before recurved ducts (Fig. 3 D). Notum 200 long (fused section 125 – 150). Gonocoxite IX small, with 3 – 4 long setae. Dorsomesal lobe with scattered microtrichia and mesal scales. Cerci 75 long by 45 – 50 wide. Tergite IX heavily setose, undivided. Pupa (n = 1, putative). 5.6 mm long. Exuviae mid-brown with darker apophyses. Cephalothorax. Cephalic area weakly creased with well developed cephalic tubercles but lacking frontal setae. Prominent scale-covered scutal hump at anterior 1 / 4. Prealar tubercle rounded. Base of thoracic horn oval, tracheal bundle densely packed, 35 x 17 µm. Abdomen (Fig. 5 B). Tergites II – VI with more extensive spinules, on II – V broad anteriorly, narrowed in midsection and posteriorly broader between apophyses; TVII – VIII with dense small spinules in anterior 1 / 3. Pleurae without spinules. Tergite II with row of 23 hooks extending 20 % of tergite width. Vortex absent. Pleural fringe only on segment VIII. Posterolateral ‘ comb’ on VIII with 11 teeth in two discrete sizes (larger lateral, smaller medial) (Fig. 4 H). Larva. Unknown / unassociated, but see Comments below and Skusella sp. ‘ Opi’.	en	Cranston, Peter S., Tang, Hongqu (2018): Skusella Freeman (Diptera: Chironomidae): new species, immature stages from Africa, Asia and Australia, and expanded distributions. Zootaxa 4450 (1): 41-65, DOI: 10.11646/zootaxa.4450.1.3
6D0F87C5011FFFDCFF3FFF0CFC25FE36.taxon	discussion	Comments. Material from Lake Opi, Nigeria collected by Landis Hare comprises 5 slides. Although all specimens belong to Skusella, the only slide (and its’ label) that shows an association is a pupal exuviae with emerged female conforming to S. freemani Harrison (see below). A second female from an emergence trap with a clearly visible bare squama, plain legs and characteristic seminal ducts conforms to S. pallidipes. The presence of two species is confirmed by two differing pupal exuviae. One has a pleural fringe extending from the posterior of segment III to VIII and is associated by rearing with the adult female identified as S. freemani. The other, an unassociated pupal exuviae from a mass rearing, has a pleural fringe restricted to segment VIII: it is treated here as belonging to S. pallidipes. Larvae cannot be allocated with certainty to either species and are treated below as Skusella sp. ‘ Opi’. It is unfortunate that in the 40 years since Hare’s pioneering work in West Africa there have been few further regional studies to reveal new life history associations.	en	Cranston, Peter S., Tang, Hongqu (2018): Skusella Freeman (Diptera: Chironomidae): new species, immature stages from Africa, Asia and Australia, and expanded distributions. Zootaxa 4450 (1): 41-65, DOI: 10.11646/zootaxa.4450.1.3
6D0F87C5011CFFDAFF3FFBE2FEA4FA5C.taxon	materials_examined	Material examined. 1 ♂, Cameroon, Adamawa [Kamerun, Adamaoŭa], 20 km S. Minim, 03. iv. 1979 (Nagel & Flacke) (ZSM). 1 Pe / ♀, Nigeria, Anambra State, Opi Lake 1 (= Ogelube Lake), 6 ° 45 ' 0 " N 7 ° 29 ' 30 " E, 10 – 13. xii. 1978, reared (Hare) (NHM). Material mapped, not examined: SOUTH AFRICA: Northern Province, Molopo Oog, Malmani Oog.	en	Cranston, Peter S., Tang, Hongqu (2018): Skusella Freeman (Diptera: Chironomidae): new species, immature stages from Africa, Asia and Australia, and expanded distributions. Zootaxa 4450 (1): 41-65, DOI: 10.11646/zootaxa.4450.1.3
6D0F87C5011CFFDAFF3FFBE2FEA4FA5C.taxon	description	Description (data from Harrison in []). Adult male (n = 1) total length 3.4 mm [5.5], wing length 1.9 mm [2.7]. Head. 4 temporals, 11 clypeals. Antennae missing [13 flagellomeres, A. R. 1.5]. Palp lengths (2 – 5): 38, 100, 100, 88 [45, 189, 201, 330]. Thorax. With single weak scutal hump. Setation: 1 L. aps, 5 – 6 [11] dc, 3 pa [3], 11 scts [6]. Wing basally infuscate with 2 broad transverse brown bands near contiguous in cells r 1, r 2 + 3, and m 1 + 2 with paler ‘ windows’ in cells r 4 + 5 and m 3 + 4. Vein setal counts R 17 [26], R 1 17 [25], R 4 + 5 13 [34]. 2 – 4 squamals. V. R. 1.05. Legs yellow except distal femur and apex of tibia (‘ knees’) on all legs brown, on forelegs distal apex of tibia also darkened. Fore and midleg tarsomeres missing. On both mid and hind tibia one comb with curved spur, the other bare. Hypopygium (Figs. 2 A – C). Tergite IX with 3 [0] anal setae. Anal point swollen, bulbous apically [rounded on holotype, more pointed on paratype]. Superior volsella strongly curved (almost hooked) medially, bearing two strong inner setae. Median volsella 30 long with simple and more taeniate setae up to 20 long (combined MV + apical taenia 50 long). Gonostylus narrowed from midpoint [moderately narrow]. Gcx 120, Gst 122, H. R. 2.8. Adult female (n = 1) (Harrison in []). Body length c. 3.5 mm [4.4]. Wing length 1.9 mm [2.6]. Head 9 – 10 temporals, 14 clypeals. Antenna 110, 70, 70, 60, 150; AR 0.42 [0.44]. Palp 2 – 5: 40, 220, 220, 310 [60, 45, 192, 207, 315]. Thorax. With double scutal hump, the anterior larger than longer, shallower posterior. Setation: 1 L. aps [1], 10 dc [15], 3 pa [3], 3 scts 10]. Wing. As in male, but fainter, with two broad separate darker bands each extending full width of wing. Vein setal counts R 23 [26], R 1 21 [27], R 4 + 5 33 + [52]. Squama damaged / missing [6]. All legs damaged but darker ‘ femoral-tibial junction (‘ knees’) visible on all legs. Genitalia as in Harrison 2002. Differences from S. pallidipes: seminal vesicles more ovoid, 90 long, maximum width 60, neck weak, less delimited from ducts, ducts initially directed posteriorly then making a complete loop anterior to opening (Fig. 3 E, as in Harrison 2002: fig. 29). Pupa (n = 1). Length 5.3 mm. Pale with darker mid-brown apophyses and golden ‘ comb’. Cephalic area with tubercles (c. 40 µm long) bearing c 30 µm frontal setae, each recessed in pit. Thoracic setation and thoracic horn structure not recognisable. Prealar tubercle present. Abdomen. Spinulation on all tergites reduced with respect to S. subvittata: on TII small triangular posteromedian area anterior to very short hook row (of 20 hooks, extending 20 % width of tergite), spinule patches on TIII – VI narrower and sparser than in S. subvittata, TVII and TVIII essentially bare (Fig. 5 C). Pedes spurii B well developed. Sternites I – VII bare, with anteromedian patch of fine spinules on VIII. ‘ Comb’ on posterolateral corner of VIII with 10 – 11 subequal-lengthed teeth very similar to that of S. subvittata (Fig. 4 H). Anal lobe fringe of 110 + / - uniserial taeniae. Larva. Unknown / unassociated, but see Skusella sp. ‘ Opi’ below.	en	Cranston, Peter S., Tang, Hongqu (2018): Skusella Freeman (Diptera: Chironomidae): new species, immature stages from Africa, Asia and Australia, and expanded distributions. Zootaxa 4450 (1): 41-65, DOI: 10.11646/zootaxa.4450.1.3
6D0F87C5011CFFDAFF3FFBE2FEA4FA5C.taxon	discussion	Comments. The adult male from Cameroon was recognized as an undescribed species of Skusella by the late Dr Friedrich Reiss of the Zoologische Staatssammlung München (Reiss in litt. to Cranston, 1989). Earlier a female reared from a pupa by Landis Hare had been identified provisionally as S. pallidipes: here we recognise this to be Skusella freemani Harrison (2002). This second African species of Skusella was described without reference to specimens now available to us. The earlier misidentification may have been due to overclearing and / or bleaching making the wing markings very faint and the characteristic leg banding indistinct. As recognized by Harrison (2002), Skusella freemani shares the patterned wing with its Afrotropical congener S. pallidipes. However it is differentiated by the banded femora and tibia, a setose squama, the shape and orientation of the superior volsella (curved and directed medially rather than posteriorly (Fig. 2 B) and tapered gonostylus (Fig. 2 A) in the male genitalia and in a looped seminal duct in the female genitalia (Fig. 3 E). The Cameroon location is some 5,000 km from the type locality in Northern Province of South Africa, and the Nigerian locality even more distant. As might be expected, some variation is seen but may derive from size as South African specimens are significantly larger than those from West Africa. The conservative approach adopted here is to consider these conspecific until further evidence is available. The associated pupa from Nigeria, described here for the first time, resembles that of S. subvittata and new forms from China in that the fringe extends from the posterior corner of pleura III through to apex of VIII. Differentiation includes the reduced spinulation on the tergites and, uniquely, an absence of spinules on the bare TVII and VIII.	en	Cranston, Peter S., Tang, Hongqu (2018): Skusella Freeman (Diptera: Chironomidae): new species, immature stages from Africa, Asia and Australia, and expanded distributions. Zootaxa 4450 (1): 41-65, DOI: 10.11646/zootaxa.4450.1.3
6D0F87C5011AFFD9FF3FFA69FEA2FA5E.taxon	description	(Figs 2 D – F, 6 A)	en	Cranston, Peter S., Tang, Hongqu (2018): Skusella Freeman (Diptera: Chironomidae): new species, immature stages from Africa, Asia and Australia, and expanded distributions. Zootaxa 4450 (1): 41-65, DOI: 10.11646/zootaxa.4450.1.3
6D0F87C5011AFFD9FF3FFA69FEA2FA5E.taxon	materials_examined	Material examined. Holotype ♂, CHINA, Guangdong Province, Guangzhou City, Conghua District, Lyutian Town, Dongkeng Reservoir, 23 ° 53.277 ' N 113 ° 55.069 ' E, 9. ix. 2015, light trap (H. Q. Tang) (EJNU). Paratypes, ♂, as holotype; Pe / ♂ (pharate male), as holotype except 3. v. 2017 (S. L. Hu & H. Q. Tang) (both ANIC). CHINA, 1 ♂, Zhejiang Province, Hangzhou City, Tianmu Mt., unnamed reservoir, 30 ° 20.02 ' N 119 ° 26.25 ' E, 800 m asl, 27. vii. 2011 (X. L. Lin); 2 ♂, CHINA, Yunnan Province, Dehong Dai & Jingpo Autonomous Prefecture, Ruili City, Mengxiu Town, Daoba country, stream in Ruili Botanic Garden, 24 ° 02 ' N 97 ° 29 ' E, 31. x. 2015 (J. Liu). Excluded. THAILAND: 1 L, Chalyaphum Prov., Amphur Chum Pae, pond, 16 ° 35 ' N 102 ° 02 ' E, 3. v. 2004, 217 m asl, # L- 660 (Vitheepradit); Pe, Kampaeng Phet Prov, Khlong Lan N. P., Nam Tok Klong Lan, 16 ° 07 ' N 99 ° 16 ' E, 8. iii. 2002, # L 272.	en	Cranston, Peter S., Tang, Hongqu (2018): Skusella Freeman (Diptera: Chironomidae): new species, immature stages from Africa, Asia and Australia, and expanded distributions. Zootaxa 4450 (1): 41-65, DOI: 10.11646/zootaxa.4450.1.3
6D0F87C5011AFFD9FF3FFA69FEA2FA5E.taxon	description	Description. Adult male (n = 6, 1 pharate). Similar to S. subvittata in colour except abdomen uniform. Legs yellow except dark brown apex of fore femur and complete fore tibia, and pale brown basal half mid and hind tibia. B. l. 3.4 – 4.0 mm, W. l. 1.6 – 1.8 mm. Fl 1 – 12 470 – 510. Fl 13 570 – 610, A. R. 1.20 – 1.25. Temporals 9 – 12 uniserial, some long; Clyp 13 – 18. Palp 2 – 5: 30 – 40, 140 – 200, 120 – 170, 205 – 258. Thorax with one distinct hump. L. aps 0 – 3. Dc 3 – 5 (small pits omitted), usually posterior pore still most posterior, accompanied by 2 tiny pits, Pa 2 – 3, Scts 6 – 8. Wing. Plain. V. R. 1.10 – 1.14. Vein setation: R 15 – 21, R 1 10 – 19, R 4 + 5 15 – 24, squamals 0 – 1. Legs. LR 1 2.26 – 2.41, LR 2 0.67 – 0.69, LR 3 0.85 – 0.95. BV 1 1.51 – 1.56, BV 2 3.20 – 3.37, BV 3 2.34 – 2.44. SV 1 1.18 – 1.21, SV 2 3.24 – 3.38, SV 3 2.44 – 2.68. Hypopygium (Fig. 2 D – F). Tergite IX with 2 – 4 median setae. Anal point slightly spatulate subapically. Superior volsella (Fig. 2 E) ca. 50 µm, with 3 – 4 lateral setae, 2 inner setae widely separated (10 – 15 µm), arising from distinct tubercle, 1 outer seta nearly always present (at least one side). Median volsella (Fig. 2 F) stem ca. 35 µm, with ca. 40 split setae. Pupa (n = 1 – 2). Body length 4.95 mm. Exuviae pale grey with brown apophyses. Cephalothorax. Cephalic tubercle low and weak. Frontal setae 50 µm, shorter than distance between setae. Thorax clearly rugose at scutal hump, otherwise smooth. Prealar tubercle seemingly absent, area damaged in dissection of pharate adult. Abdomen (Fig. 6 A). Similar to S. subvittata, with more extensive tergal spinulation. Pleural fringe slightly longer on IV – VII (90 – 100 µm) than others (50 – 60 µm), on segment VIII slightly different to other segment, anterior ca. 75 µm, posterior ca. 150 µm. Hook row of 26 – 31 hooks, extending 27 – 30 % of segment width. Posterolateral corner of VIII (‘ comb’) with 9 – 12 teeth of variable length. Anal lobe with 116 – 118 taeniae. Larva. Material from Thailand may be conspecific to this species, but status will need molecular association. Female unknown.	en	Cranston, Peter S., Tang, Hongqu (2018): Skusella Freeman (Diptera: Chironomidae): new species, immature stages from Africa, Asia and Australia, and expanded distributions. Zootaxa 4450 (1): 41-65, DOI: 10.11646/zootaxa.4450.1.3
6D0F87C5011AFFD9FF3FFA69FEA2FA5E.taxon	etymology	Etymology (Derivatio nominis). The species epithet honors the junior author’s mother, Siling Hu, who helped collect the pharate material during a camping field trip.	en	Cranston, Peter S., Tang, Hongqu (2018): Skusella Freeman (Diptera: Chironomidae): new species, immature stages from Africa, Asia and Australia, and expanded distributions. Zootaxa 4450 (1): 41-65, DOI: 10.11646/zootaxa.4450.1.3
6D0F87C5011AFFD9FF3FFA69FEA2FA5E.taxon	discussion	Comments. Skusella silingae resembles the type species, S. subvittata, in the known life stages. The wing is unmarked, the anal point is more spatulate distally with a rounded apex and the superior volsella has 3 – 4 lateral setae, of which 1 outer seta always is present, the apex tapering more abruptly (Fig. 2 E). The pupa has a pleural fringe extending from the posterior of TIII to VIII, spinulation on tergites II – IV not extending laterally beyond adhesion marks, and with at least 25 hooklets on TII, row extending more than 1 / 4 of segment width. Thai material is placed here as an atypical form of S. silingae since the pupal hook row has 31 hooklets, occupying ca. 30 % of segment width, both measures greater than in the Chinese paratype. The type locality is a relatively clean mountain reservoir. The water temperature ranges from 13.8 ° C to 25.1 ° C, dissolved oxygen 6.0 – 7.2 mg / l, pH 6.7 – 7.8, total nitrogen 0.1 – 0.6 mg / l, total phosphorus 0.04 – 0.10 mg / l, and COD 2.4 – 15.2 mg / l.	en	Cranston, Peter S., Tang, Hongqu (2018): Skusella Freeman (Diptera: Chironomidae): new species, immature stages from Africa, Asia and Australia, and expanded distributions. Zootaxa 4450 (1): 41-65, DOI: 10.11646/zootaxa.4450.1.3
6D0F87C50119FFC6FF3FF9ACFC03FEA6.taxon	materials_examined	Material examined. Pe (♀), slide mounted in Canada balsam, CHINA, Hainan [Jianfengling NNR, drift sample from stream at approx. 1000 m elevation], 23. v. 1980 (Fittkau) (ZSM).	en	Cranston, Peter S., Tang, Hongqu (2018): Skusella Freeman (Diptera: Chironomidae): new species, immature stages from Africa, Asia and Australia, and expanded distributions. Zootaxa 4450 (1): 41-65, DOI: 10.11646/zootaxa.4450.1.3
6D0F87C50119FFC6FF3FF9ACFC03FEA6.taxon	description	Description. Pupa (n = 1). 5.2 mm long. Exuviae pale brown with darker brown abdominal apophyses. Cephalothorax. Cephalic area smooth with minute frontal setae arising from pale area of cuticle. Thorax smooth excepting oval patch of rounded scales in mid-thorax that may be associated with weak scutal ‘ hump’. Base of thoracic horn elongate-oval, tracheal bundle sparsely packed, 17 x 19 µm. Without prealar tubercle. Abdomen. Tergites II – VI with extensive spinulation, anteriorly filling area between apophyses and anteriorly extending to pleurae (Fig. 4 F). Tergite II with row of 23 hooks extending 25 % of tergite width. Pedes spurii B moderately developed, tapering, projecting ventrally rather than laterally (perhaps according to slide preparation). Pleural fringe on segment VIII only. Posterolateral ‘ comb’ on VIII with 11 teeth diminishing in size from lateral to medial (Fig. 4 J).	en	Cranston, Peter S., Tang, Hongqu (2018): Skusella Freeman (Diptera: Chironomidae): new species, immature stages from Africa, Asia and Australia, and expanded distributions. Zootaxa 4450 (1): 41-65, DOI: 10.11646/zootaxa.4450.1.3
6D0F87C50119FFC6FF3FF9ACFC03FEA6.taxon	discussion	Comments. This unreared pupal exuviae from Jianfengling, Hainan, shares uniquely with Afrotropical S. pallidipes a pleural fringe restricted only to segment VIII. The pupal exuviae can be distinguished principally by the tergal spinulation on III – VII extending clearly onto the pleurae (Fig. 4 F). In contrast in S. pallidipes all pleurae lack spinules in a much more restricted spinulation pattern (Fig. 5 B).	en	Cranston, Peter S., Tang, Hongqu (2018): Skusella Freeman (Diptera: Chironomidae): new species, immature stages from Africa, Asia and Australia, and expanded distributions. Zootaxa 4450 (1): 41-65, DOI: 10.11646/zootaxa.4450.1.3
6D0F87C50106FFC6FF3FFE31FBB5FBF8.taxon	materials_examined	Material examined. 5 Pe, CHINA, Hainan Province, Qiongzhong County, Limuling NNR, 19 ° 10.755 ' N 109 ° 45.074 ' E, 11. ii. 2015 (H. Q. Tang) (EJNU, 2 (1 slide) to ANIC); 3 Pe, CHINA, Hainan Province, Lingshui Li Autonomous County, Diaoluo National Forest Park, 18 ° 43.606 ' N 109 ° 52.051 ' E, 26. iv. 2012 (H. Q. Tang); 1 Pe, CHINA, Hainan Province, Changjiang Li Autonomous County, Bawangling NNR, 19 ° 06.752 ' N 109 ° 11.980 ' E, 30. iv. 2012 (H. Q. Tang); 1 Pe, CHINA, Hainan Province, Wanning City, 3 km away from Martyrs Memorial Park of Liulianling, 18 ° 58.825 ' N 110 ° 25.348 ' E, 09. ii. 2015 (H. Q. Tang).	en	Cranston, Peter S., Tang, Hongqu (2018): Skusella Freeman (Diptera: Chironomidae): new species, immature stages from Africa, Asia and Australia, and expanded distributions. Zootaxa 4450 (1): 41-65, DOI: 10.11646/zootaxa.4450.1.3
6D0F87C50106FFC6FF3FFE31FBB5FBF8.taxon	description	Description. Pupa (n = 5). Body length 5.2 – 5.6 mm. Exuviae pale grey with brown apophyses. Cephalothorax. Cephalic tubercle absent. Frontal setae 40 – 50 µm, half the length of distance between setae (Fig. 4 A). Thorax clearly rugose at scutual hump, otherwise smooth (Fig. 4 B). Prealar tubercle absent. Abdomen (Fig. 6 B). Similar to S. silingae but with spinulae on anterior of each tergite extending more laterad including to onto anterior pleurae. Hook row of 24 – 27 hooks, extending 23 – 28 % of segment width. Posterolateral corner of VIII (‘ comb’) with 9 – 12 teeth of variable length. Anal lobe with 130 – 140 taeniae.	en	Cranston, Peter S., Tang, Hongqu (2018): Skusella Freeman (Diptera: Chironomidae): new species, immature stages from Africa, Asia and Australia, and expanded distributions. Zootaxa 4450 (1): 41-65, DOI: 10.11646/zootaxa.4450.1.3
6D0F87C50106FFC6FF3FFE31FBB5FBF8.taxon	discussion	Comments. The pupal exuviae resemble those of S. subvittata and S. silingae but lacks a cephalic tubercle and has more extensive anterior tergal spinulation. From differs from S. sp. ‘ Sanyatang’ in the spinulosity on the pleurae being restricted to small areas on segments V – VII and spinules are lacking on sternite I. Exuviae were collected from some near pristine mountain streams and ponds.	en	Cranston, Peter S., Tang, Hongqu (2018): Skusella Freeman (Diptera: Chironomidae): new species, immature stages from Africa, Asia and Australia, and expanded distributions. Zootaxa 4450 (1): 41-65, DOI: 10.11646/zootaxa.4450.1.3
6D0F87C50106FFC6FF3FFB1BFE1FF90A.taxon	materials_examined	Material examined. 1 Pe (♀), CHINA, Guangdong Province, Guangzhou City, Conghua District, Sanyatang Hydropower Station, 23 ° 44.468 ′ N 113 ° 48.216 ′ E, 28. vii. 2014 (H. Q. Tang) (ANIC). 1 Pe (♂), CHINA, Guangdong Province, Guangzhou City, Zengcheng, pool area of Lan stream, a dam at Shuimei Country, 23 ° 21.318 ′ N 113 ° 58.070 ′ E, 20. xii. 2017 (H. Q. Tang) (EJNU),	en	Cranston, Peter S., Tang, Hongqu (2018): Skusella Freeman (Diptera: Chironomidae): new species, immature stages from Africa, Asia and Australia, and expanded distributions. Zootaxa 4450 (1): 41-65, DOI: 10.11646/zootaxa.4450.1.3
6D0F87C50106FFC6FF3FFB1BFE1FF90A.taxon	description	Description. Pupa (n = 2). Body length 4.1 – 4.4 mm. Exuviae pale brown with brown abdominal apophyses. Cephalothorax. Cephalic tubercle absent. Frontal setae 40 µm. Thorax granulate only in middle, with 8 – 10 rounded small beads. Prealar tubercle absent. Abdomen (Fig. 6 C). Spinulation extensive on II – VII, extending onto most pleurae lateral to adhesion marks. Hook row of 18 – 24 hooks, 18 – 22 % of width of segment II. Comb with 8 – 12 teeth. Sternite 1 (Fig. 4 E) with anteromedian and anterolateral weak spinule patches.	en	Cranston, Peter S., Tang, Hongqu (2018): Skusella Freeman (Diptera: Chironomidae): new species, immature stages from Africa, Asia and Australia, and expanded distributions. Zootaxa 4450 (1): 41-65, DOI: 10.11646/zootaxa.4450.1.3
6D0F87C50106FFC6FF3FFB1BFE1FF90A.taxon	discussion	Comments. Skusella sp. ‘ Sanyatang’ pupal exuviae generally resemble those of S. subvittata and S. silingae but differ in more extensive spinulation of tergites II to IV and with these pleurae being widely spinulose (Fig. 6 C). From S. sp. ‘ Limuling’ this taxon is separated by more extensive pleural spinulation and the presence of fine spinules on sternite I.	en	Cranston, Peter S., Tang, Hongqu (2018): Skusella Freeman (Diptera: Chironomidae): new species, immature stages from Africa, Asia and Australia, and expanded distributions. Zootaxa 4450 (1): 41-65, DOI: 10.11646/zootaxa.4450.1.3
6D0F87C50106FFC2FF3FF8D4FC7CFE83.taxon	materials_examined	Material examined. NIGERIA, Anambra State, 2 L (3 rd i., 4 th i), Opi Lake ‘ A’ [= Ogelube Lake], 6 ° 45 ' 0 " N 7 ° 29 ' 30 " E, 1. iii. 1977 (Hare) (NHM); same except 1 L (2 i), Opi Lake ‘ D’, 15. v. 1980.	en	Cranston, Peter S., Tang, Hongqu (2018): Skusella Freeman (Diptera: Chironomidae): new species, immature stages from Africa, Asia and Australia, and expanded distributions. Zootaxa 4450 (1): 41-65, DOI: 10.11646/zootaxa.4450.1.3
6D0F87C50106FFC2FF3FF8D4FC7CFE83.taxon	description	Description. Larva 4 i (n = 1). Head pale yellow with slightly darker golden-brown occipital margin, mandibles and mentum. Body length (extended exuviae) c. 10 mm, head capsule 500 µm, postmentum 150 µm. Antenna (Fig. 7 C) segment lengths 115: 18: 45: 30: 10, 5; AR 1.06. Lauterborn organs 10 long; blade c. 60 long, extending to mid-segment 3. Labroepipharyngeal region as in S. subvittata, Premandible 65 long. Mandible 105. Mentum (Fig. 7 H) 90 wide; ventromental plates 100 wide, separated medially by 50, VmPR 2.2. Abdomen. Anterior parapod claws mid-yellow, simple, narrow, gently curved. Procercus pale, squat, 20 wide, 12 high, bearing 2 fine lateral seta, respectively 15 and 25 long; apically with 5 pale yellow setae 250 – 300 long. Posterior parapod claws pale yellow, simple, finely curved. Larva 3 i (n = 1) Head pale. Head capsule 250, submentum 105. Antenna 55, 8, 27, 17, 7, 4; A. R. 0.9. Premandible 65. Mandible 70. Mentum 53 wide; ventromental plates 65, separated medially by 45, VmPR 2.6. Larva 2 i (n = 1). Head pale, body red (stained?). Body length 1.7 mm. Head capsule 150, submentum 60. Antenna 30, 6, 15, 9, 4.5, 3; A. R. 0.9. Premandible 30. Mandible 70. Mentum 37 wide; ventromental plates 33, separated medially by 20, VmPR 2.6.	en	Cranston, Peter S., Tang, Hongqu (2018): Skusella Freeman (Diptera: Chironomidae): new species, immature stages from Africa, Asia and Australia, and expanded distributions. Zootaxa 4450 (1): 41-65, DOI: 10.11646/zootaxa.4450.1.3
6D0F87C50106FFC2FF3FF8D4FC7CFE83.taxon	discussion	Comments. Material from Landis Hare from Nigeria include larval head capsules of 3 rd and 4 th instars. Additionally a larval exuviae under 3 coverslips on one slide is seemingly associated with the 4 th instar head capsule. A second slide labelled as ‘ 2 i’, contains an unreared larva, of appropriate size for a second instar judging from application of Dyar’s rule (% increment increase in sclerotized structure such as mandible from 3 rd to 4 th instar) of 1.66 between the associated 3 rd and 4 th instars. No larva can be associated specifically with either S. pallidipes or S. freemani (see above). However these unallocated larvae demonstrate that larvae clearly congeneric with those of the Australian genotype (S. subvittata) were present at a location in West Africa where two identified species of Skusella occur. Details of the larval habitat, a dilute water lake in the transition between guinea savannah and lowland moist forest in south-eastern Nigeria can be found in Hare & Carter (1984, 1987).	en	Cranston, Peter S., Tang, Hongqu (2018): Skusella Freeman (Diptera: Chironomidae): new species, immature stages from Africa, Asia and Australia, and expanded distributions. Zootaxa 4450 (1): 41-65, DOI: 10.11646/zootaxa.4450.1.3
