Papuanatula (Papuanatula) batanlenos Kaltenbach, Kovács & Gattolliat sp. nov. Figs 15, 16, 17, 18, 19 Type material. Holotype. Indonesia • larva; West Papua, Batanta Island, Kalijakut River; 00°52'27"S, 130°37'52"E; 420 m; 09. ii. 2024; leg. T. Kovács; on slide; GBIFCH 01221820; 2024-12, EPHTYP-11; MM. Paratypes. • 14 larvae; same data as holotype; • 2 larvae on slides; GBIFCH 00975879, GBIFCH 01221821 (gonostyli), GBIFCH 01221824; MZL; • 12 larvae in alcohol; GBIFCH 00975880, GBIFCH 00975881, GBIFCH 00975878, GBIFCH 01581951; MZL • 3 subimagos; same data as holotype; at light, in alcohol; GBIFCH 00975863 (♂, ♀), GBIFCH 00975864 (♀); MZL • 2 larvae; West Papua, Batanta Island, Kalijakut River; between 00°53'39"S, 130°38'30"E and 00°53'03"S, 130°38'13"E; 52 m – 182 m; 08. ii. 2024; leg. T. Kovács; in alcohol; GBIFCH 00975874, GBIFCH 00975875; MZL • 4 larvae; West Papua, Batanta Island, valley of Waridor River; 00°51'52"S, 130°32'26"E; 55 m; 19. i. 2014; leg. T. Kovács, R. Horváth, P. Juhász; 1 on slide; GBIFCH 00975994; 3 in alcohol; GBIFCH 00975995, GBIFCH 00975996; MZL • 1 ♂ imago, 3 subimagos; West Papua, Batanta Island, valley of Kalijakut River; 00°52'49"S, 130°38'05"E; 232 m; 19. ii. 2020; at light; leg. T. Kovács, R. Horváth, P. Juhász, K. Sauyai, R. Sauyai; 1 ♂ imago on slide, thorax in alcohol; GBIFCH 01221780; GBIFCH 00975967; 3 subimagos in alcohol; GBIFCH 00975965 (♂, ♀), GBIFCH 00975966 (♂); MZL • 1 imago, 2 subimagos; West Papua, Batanta Island, valley of Kalijakut River; 00°53'03"S, 130°38'13"E; 182 m; 15. ii. 2023; at light; leg. T. Kovács, R. Horváth, P. Juhász, K. Sauyai, R. Sauyai; 1 ♂ imago on slide, thorax in alcohol; GBIFCH 01221779, GBIFCH 00975970; 1 subimago on slide; GBIFCH 00975968 (♂); 1 subimago in alcohol; GBIFCH 00975969 (♀); MZL • 1 ♂ imago; West Papua, Batanta Island, valley of Tanjung Lampu River; 00°53'43"S, 130°36'39"E; 175 m; 21. ii. 2018; at light; leg. T. Kovács, R. Horváth, P. Juhász, K. Sauyai, R. Sauyai; on slide, thorax in alcohol; GBIFCH 01221776, GBIFCH 00975964; MZL • 5 subimagos; West Papua, Batanta Island, valley of Wailebet Stream; 00°53'22"S, 130°39'07"E; 150 m; 19. ii. 2018; at light; leg. T. Kovács, R. Horváth, P. Juhász, K. Sauyai, R. Sauyai; 4 in alcohol; GBIFCH 00975961 (1 ♂, 2 ♀), GBIFCH 00975962 (♀); 1 on slide; GBIFCH 00975963 (♀); MZL • 3 ♀ subimagos; West Papua, Batanta Island, valley of Wailebet Stream, waterfall; 00°53'07"S, 130°39'00"E; 285 m; 28. ii. 2017; at light; leg. T. Kovács, R. Horváth, P. Juhász, K. Sauyai, R. Sauyai; in alcohol; GBIFCH 00975959, GBIFCH 00975960; MZL • 15 larvae; West Papua, Batanta Island, Waridor River (shallow, rocky, fast-flowing); 00°52'06"S, 130°31'30"E; 32 m; 14. ii. 2025; leg. T. Kovács; 6 in alcohol; GBIFCH 01582001; MZL; 9 in alcohol; 2025-12, EPHTYP-12; MM • 10 larvae; Indonesia, West Papua, Batanta Island, Kalijakut River; 00°53'03"S, 130°38'13"E; 182 m; 19. ii. 2025; leg. T. Kovács; in alcohol; 2025-17, EPHTYP-13; MM • 15 larvae; West Papua, Batanta Island, Kalijakut River; 00°52'27"S, 130°37'52"E; 420 m; 20. ii. 2025; leg. T. Kovács; in alcohol; 2025-18.a, EPHTYP-14; MM • 3 larvae; West Papua, Batanta Island, right side stream of Kalijakut River; 00°52'16"S, 130°37'45.5"E; 545 m; 21. ii. 2025; leg. T. Kovács; in alcohol; 2025-19, EPHTYP-15; MM. Diagnosis. Larva. The following combination of characters distinguishes P. batanlenos sp. nov. from other species of Papuanatula s. str.: body dorsally without row of long, fine, simple setae along midline; body dorsally without protuberances; abdomen dorsally grey-brown, abdominal terga IV, VII, and VIII with trough-like, dark grey-brown marking, abdominal terga III, V, and VI with dark grey-brown, oblique, lateral streaks; femur with irregular, shoe-shaped marking in large, basal blank; paracercus with 14–16 segments; paraproct without posterior prolongation. Description. Larva (Figs 15 – 17). Body length 3.5–4.0 mm, cerci ~ 1.6 × body length. Cuticular coloration (Figs 15 a – c, 17 e). Head, thorax and abdomen dorsally grey-brown, thorax darker; abdominal terga IV, VII, and VIII with trough-like, dark grey-brown marking; abdominal terga III, V, and VI with dark grey-brown, oblique, lateral streaks (dorsal abdominal pattern slightly varying). Head, thorax and abdomen ventrally grey-brown. Legs grey-brown, with large blank in basal area. Caudalii yellow-brown. Hypodermal coloration (Fig. 15 a – c). Abdominal terga I – IX with narrow, dark brown, transverse band along posterior margin. Femur anteriorly with dark brown, irregular, shoe-shaped marking in basal ½ (somewhat variable in shape; sometimes poorly developed); posteriorly with dark brown streak distally close to outer margin. Head. Antenna (Fig. 16 j). Length ~ 2.5 × head length. As typical for the subgenus. Developing turbinate eyes in last instar male larva (Fig. 15 a) large, round, nearly touching each other in the middle. Labrum (Fig. 16 a, b). Length 0.5 × maximum width, laterally convex. Dorsal, sub-marginal arc with ~ 15 feathered setae. Right mandible (Fig. 16 d). Margin between prostheca and mola straight, with few minute denticles. Otherwise, as typical for subgenus. Left mandible (Fig. 16 c). Margin between prostheca and mola straight, with few minute denticles. Subtriangular process with undulating margins, outer margin with denticles. Otherwise, as typical for subgenus. Hypopharynx (Fig. 16 f). Medial tuft of spine-like setae laterally denser, giving the impression of a pair of tufts. Otherwise, as typical for the genus. Maxilla (Fig. 16 g). Maxillary palp slightly longer than galea-lacinia; palp segment II ~ 1.2 × length of segment I. Otherwise, as typical for the genus. Labium (Fig. 16 e). Paraglossa with two spine-like setae on inner, distolateral margin. Labial palp with segment I ~ 0.8 × length of segments II and III combined. Segment II without distomedial protuberance, dorsally with row of four spine-like setae near outer, distolateral margin. Segment III broad pentagonal, pointed, 0.8 × length of segment II. Otherwise, as typical for the genus. Thorax. Sterna without protuberances. Terga without protuberances. Legs (Fig. 17 a – d). Ratio of leg segments: fore leg 1.0: 1.0: 0.2: 0.1, middle leg 1.0: 1.0: 0.2: 0.1 and hind leg 1.1: 1.0: 0.2: 0.1. Femur. Length 3 × maximum width. Many short, spine-like setae along ventral margin. Claw with one row of six or seven denticles, distalmost denticle with distance to other denticles; one or two posterior setae. Otherwise, as typical for subgenus. Abdomen. Terga (Fig. 17 f). Abdominal terga without protuberances. Posterior margin of terga: I smooth, without denticles; II – IX with triangular, pointed denticles. Tergalii (Fig. 16 h, i). Ovoid, tracheation rather well developed; margin with minute serration and short, fine, simple setae. Tergalius II as long as abdominal terga III, IV, and ¼ V combined, tergalius IV as long as terga V, VI and ⅓ VII combined, tergalius VII as long as terga VIII and ⅓ IX combined. Paraproct (Fig. 17 g). Posterior margin without prolongation, smooth, without denticles. Caudalii (Fig. 17 h, i). Cerci without swimming setae. Paracercus with 14–16 segments. Pose of subimaginal gonostyli under larval cuticle (Fig. 16 k). Segment III conical. Otherwise, as typical for the subgenus. Subimago (Fig. 19 b – f). Body length 4.2–4.5 mm. Turbinate eyes of male ochre. Thorax brown. Wing membrane colourless, veins pale yellow-brown, base of wings brown, microtrichia pale yellow-brown. Legs very pale brown; coxa with hypodermal, dark brown fleck; femur anteriorly with hypodermal, shoe-shaped, dark brown marking in basal ½; femur posteriorly with hypodermal, dark brown distal streak and fleck in basal ½. Abdomen yellow-brown; male with hypodermal, dark brown, trough-shaped marking on terga III, IV, VII, and VIII, and sublateral, oblique, dark brown streaks on terga II, V and VI; female with same hypodermal markings, but more diffuse; all terga of both sexes with hypodermal, narrow, transvers, dark brown band on posterior margin. Cerci colourless. Texture (Fig. 19 e, f). On all legs of male and female subimagos, terminal tarsomere covered with pointed microlepides; other tarsomeres covered mostly with blunt microlepides, with pointed microlepides near apex. Imago, male (Figs 18 a – d, 19 a). Body length ~ 3.8 mm. Head and antennae pale brown. Turbinate eyes pale brown, wide, with facetted surfaces round. Thorax pale brown. Fore wing with membrane colourless, base of wings brownish. Pterostigma with three or four oblique crossveins, basal two crossveins nearly complete. Legs light brownish; coxa with hypodermal, dark brown fleck; femur anteriorly with hypodermal, shoe-shaped, dark brown marking in basal ½; femur posteriorly with hypodermal, dark brown distal streak and fleck in basal ½. Abdominal segments I and VII – X pale brown, II – VI transparent, giving a blueish impression; with hypodermal, dark brown, trough-shaped marking on terga III, IV, VII, and VIII, and sublateral, oblique, dark brown streaks on terga II, V, and VI. Cerci colourless. Genitalia (Fig. 18 d). Unistyliger slightly widened apically, with lateral margin slightly concave. Gonostylus with segment I conical and separated from segment II by concavity. Segment II slightly wider in distal ½, bent inward. Terminal segment III narrower than segment II, ovoid, length ~ 1.5 × maximal width. Penial bridge with poorly expressed membranous projection between unistyligers. Imago, female. Unknown. Egg. Unknown. Eggs extracted from female subimagos are undeveloped. Biological aspects. It is a common species in most riverine habitat types in Batanta, living in the northern and southern watercourses of the island: Kalijakut (Fig. 29 a, d (type locality), f), Tanjung Lampu and Waridor (Fig. 29 e) Rivers, and Wailebet Stream. It is found from small, high-falling mountain tributaries to the lower, flat sections of the largest rivers at altitudes between 32 m and 545 m. Apart from the extr
Papuanatula (Papuafiliola) horvathrobi Kovács, Kaltenbach & Gattolliat sp. nov. Figs 20, 21, 22 Type material. Holotype. Indonesia • larva; West Papua, Batanta Island, Waibin River; between 00°49'21"S, 130°45'57"E and 00°50'02"S, 130°45'25"E; 20 m – 45 m; 05. ii. 2024; leg. T. Kovács; on slide; GBIFCH 00975876; 2024-9, EPHTYP-16; MM. Paratypes. 1 larva; same data as holotype; on slide; GBIFCH 00975913; MZL • 1 larva; West Papua, Batanta Island, Warai Stream; between 00°50'51"S, 130°35'14"E and 00°51'12"S, 130°35'20"E; 225 m – 315 m; 21. ii. 2024; leg. T. Kovács; on slide; GBIFCH 00975877; MZL • West Papua, Batanta Island, Waridor River (shallow, rocky, fast-flowing); 00°52'06"S, 130°31'30"E; 32 m; 14. ii. 2025; leg. T. Kovács; 4 in alcohol; GBIFCH 01582003; MZL; 12 in alcohol; 2025-12, EPHTYP-17; MM. Diagnosis. Larva. The following combination of characters distinguishes P. horvathrobi sp. nov. from other species of Papuanatula (Papuafiliola): thorax and abdomen dorsally without protuberances; femur with large blank in posterior ⅔; femur and abdomen without brown hypodermal maculae; tergalii oblong; paracercus with 15 segments. Description. Larva (Fig. 20 – 22). Body length ~ 3.0 mm, cerci ~ 1.4 × body length. Cuticular coloration (Figs 20 a, b, 22 a). Head, thorax, and abdomen dorsally brown. Head, thorax and abdomen ventrally beige. Legs brown, femur with large blank in posterior ⅔, and distally with small blank area. Caudalii light brown. Hypodermal coloration (Fig. 20 a, b). Head, thorax, abdomen, and legs without hypodermal markings. Head. Antenna. Length ~ 2.5 × head length. Otherwise, as typical for the subgenus. Developing turbinate eyes in last instar male larva (Figs 20 a, 21 j) large, roundish, with some distance to each other. Labrum (Fig. 21 a, b). Length 0.6 × maximal width. With reduced, submarginal arc of three long, simple setae. Otherwise, as typical for subgenus. Right mandible (Fig. 21 d). Incisor with five denticles; kinetodontium with three denticles, inner lateral margin with row of small denticles, becoming smaller and finer toward base, and with row of short, fine setae; margin between prostheca and mola slightly convex, with few minute denticles. Otherwise, as typical for subgenus. Left mandible (Fig. 21 c). Incisor with five denticles, kinetodontium with three denticles; margin between prostheca and subtriangular process straight, with minute denticles. Otherwise, as typical for subgenus. Hypopharynx (Fig. 21 f) with well-developed tuft of long, spine-like setae. Otherwise, as typical for the genus. Maxilla (Fig. 21 g). Maxillary palp slightly longer than galea-lacinia; segment I 1.4 × length of segment II. Otherwise, as typical for the genus. Labium (Fig. 21 e) Paraglossa dorsally with three spine-like setae near inner, distolateral margin. Labial palp with segment I 0.7 × length of segments II and III combined. Segment II with narrow thumb-like distomedial protuberance, slightly bent distad, dorsally with one spine-like seta near outer, distolateral margin. Segment III oblique conical, apically rounded, approx. as long as segment II. Otherwise, as typical for the genus. Thorax. Sterna without protuberances. Terga. Without long setae on midline; without protuberance. Legs (Fig. 22 a – d). Ratio of leg segments: fore leg 1.1: 1.0: 0.3: 0.1, middle leg 1.2: 1.0: 0.3: 0.1 and hind leg 1.5: 1.0: 0.4: 0.2. Femur. Length ~ 3 × maximum width. Surface of brown areas with minute denticles. Tarsus. With regular row of medium, apically rounded setae along outer margin. Claw with one row of 11 denticles; one posterior seta. Otherwise, as typical for subgenus. Abdomen. Terga (Fig. 22 e, f, j). Abdominal terga without long setae on midline; abdominal terga without median or submedian elevations or protuberances; surface with minute denticles. Posterior margins of abdominal terga: I smooth, without denticles, II – IX with very small, triangular denticles. Tergalii (Fig. 21 h, i). Present on terga II – VII, oblong; tracheation well-developed, pigmentation not reaching margins. Each tergalius with anal rib longer than costal rib; ribs with minute, irregularly situated denticles on dorsal side. Tergalius II as long as terga III and ⅔ IV combined, tergalius IV as long as terga V and ⅔ VI combined, tergalius VII as long as terga VIII and ⅔ IX combined. Paraproct (Fig. 22 g) without posterior prolongation. Posterior margin with small denticles. Caudalii (Fig. 22 h, i). Cerci without swimming setae. Paracercus with 15 segments. Pose of subimaginal gonostyli under larval cuticle unknown. Subimago. Unknown. Imago. Unknown. Egg. Unknown. Biological aspects. The species is known from three watercourses in the northern part of Batanta (Waibin River (type locality), Waridor River, and Warai Stream), at altitudes from 20– 315 m. The largest numbers were collected at a ford of the Waridor River at an altitude of 32 m (Fig. 29 e), where P. batanlenos sp. nov., was also present in large numbers. Etymology. The new species is dedicated to Róbert Horváth, a Hungarian ornithologist, who initiated and organized the Hungarian research program on Biodiversity in Batanta in 2010. Since then, he gave continuous support in various aspects of the ten research trips conducted to date. Distribution. New Guinea, Batanta Island (Fig. 30).
Labiobaetis lahu sp. nov. Figs 4, 5, 6 Type material. Holotype. Thailand • larva; Tak Prov., Mueang Distr., Klong Lan Sang; 16°46'53"N, 99°01'16"E; 253 m; 25.xii.2017; leg. C. Suttinun; on slide; GBIFCH 00596154; VMCMU. Paratypes. 2 larvae; same data as holotype; on slides; GBIFCH 01223072, GBIFCH 01223073; MZL • 2 larvae; Tak Prov., Mueang Distr., Oum Yom; 06°48'16"N, 99°01'19"E; 249 m; 26.xii.2017; leg. C. Suttinun; 1 on slide; GBIFCH 01223071; 1 in alcohol; GBIFCH 00763789; VMCMU • 8 larvae; Loei Prov., Wang Saphung Distr., Loei River; 17°18'04"N, 101°46'33"E; 256 m; 18.xii.2018; leg. C. Suttinun; 1 on slide; GBIFCH 00607175; 7 in alcohol; GBIFCH 00763787, GBIFCH 00763790; VMCMU • 22 larvae; Chiang Mai Prov., Chiang Dao Distr., Kaeng Pantao; 19°17'02"N, 98°58'05"E; 403 m; 21.xi.2018 and 05.xi.2023; leg. C. Suttinun; 1 on slide; GBIFCH 00607176; MZL; 21 in alcohol; GBIFCH 00980856, GBIFCH 00980858, GBIFCH 00763791, GBIFCH 00763786, GBIFCH 00763871, GBIFCH 00763874; VMCMU • 63 larvae; Tak Prov., Mae Sot Distr., Moei River; 16°42'04"N, 98°30'41"E; 194 m; 27.xii.2017; leg. C. Suttinun; in alcohol; GBIFCH 00829295, GBIFCH 00763785, GBIFCH 00763788; VMCMU; GBIFCH 00763851, GBIFCH 00763873; MZL • 23 larvae; Ratchaburi Prov., Suan Phueng Distr., Pha Wo Tai; 13°30'57"N, 99°20'40"E; 118 m; 25.xi.2018; leg. C. Suttinun; in alcohol; GBIFCH 00980851, GBIFCH 00763807; VMCMU. Diagnosis. Larva. Following combination of characters differentiate L. lahu sp. nov. from other species of the group batakorum: thorax and abdomen dorsally dark grey with yellowish pattern as in Fig. 4 a, especially yellowish oval markings on terga III and V; legs with reddish-brown spot distally on femur; hind protoptera present, well developed. Description. Larva (Figs 4 – 6). Body length 4.2–5.0 mm. Cerci ~ 0.4 body length, paracercus ~ 0.3 × body length. Antenna: ~ 2 × as long as head length. Colouration (Fig. 4 a – c). Thorax and abdomen dorsally dark grey with distinct yellowish markings as in Fig. 1 a; abdominal segments V and X brighter. Fore protoptera yellowish and grey. Thorax and abdomen ventrally whitish-yellow; abdominal segments II – V with dark grey, arched marking; segments VI – IX dark grey. Legs off-white to yellowish, femur medially and basally with grey markings, distally with distinct reddish-brown spot, tibia and tarsus distally grey. Caudalii whitish-yellow, dark brown section in distomedial part. Antenna (Fig. 6 i) with scape and pedicel sub cylindrical, distolateral process at scape well developed. Labrum (Fig. 5 a, b). Sub-rectangular, length 0.7 × maximum width. Distal margin with medial emargination and small process. Dorsally with medium, fine, simple setae scattered over surface; pair of submedian setae, and submarginal arc of four or five long, simple setae on each side, 1 st and 2 nd setae closely together. Ventrally with marginal row of setae composed of lateral and anterolateral long, feathered setae and medial long, bifid setae. Right mandible (Fig. 5 i). Incisor and kinetodontium fused. Incisor with four denticles; kinetodontium with four denticles, inner margin of innermost denticle with row of thin setae. Prostheca robust, apically denticulate. Margin between prostheca and mola slightly convex, smooth. Tuft of setae on proximal corner of mola present. Left mandible (Fig. 5 h). Incisor and kinetodontium fused. Incisor with four denticles, kinetodontium with three denticles. Prostheca robust, apicolaterally with small denticles and comb-shaped structure. Margin between prostheca and mola slightly convex, smooth. Tuft of setae on proximal corner of mola absent. Both mandibles with lateral margins almost straight. Hypopharynx and superlinguae (Fig. 5 e). Lingua longer than superlinguae. Lingua longer than broad; medial tuft of stout setae well developed, short; in distal 1 / 2 expanded. Superlinguae with lateral margins rounded; fine, long, simple setae along distal margin. Maxilla (Fig. 5 f, g). Galea-lacinia ventrally with two simple, apical setae below canines. Medially with one feathered, spine-like seta and four short to long, simple setae. Maxillary palp longer than length of galea-lacinia; 2 - segmented; palp segment II approx. as long as segment I; setae on maxillary palp fine, simple, scattered over surface of segments I and II; apex of last segment with distolateral excavation, apically constricted. Labium (Fig. 5 c, d). Glossa basally broad, narrowing toward apex; shorter than paraglossa; inner margin with ~ 9 spine-like seta; apex with three long, robust, apically pectinate setae; outer margin with ~ 4 spine-like setae; ventral surface with fine, simple, scattered setae. Paraglossa sub-rectangular, slightly curved inward; apex rounded; with three rows of long, robust, distally pectinate setae in apical area and one short, simple seta in anteromedial area; dorsally with five long, spine-like setae near inner margin. Labial palp with segment I ~ 0.8 × length of segments II and III combined. Segment II with broadly rounded, thumb-like, distomedial protuberance; distomedial protuberance 0.7 × width of base of segment III; dorsally with row of ~ 4 spine-like setae near outer margin. Segment III conical; length ~ 0.8 × maximal width; ventrally covered with short, spine-like, simple setae and short, fine, simple setae. Hind protoptera (Fig. 6 e) present, well developed. Legs (Fig. 6 a – d). Ratio of foreleg segments 1.4: 1.0: 0.6: 0.2, middle leg 1.3: 1.0: 0.6: 0.2, hind leg 1.2: 1.0: 0.7: 0.2. Femur. Femur length ~ 3 × maximum width. Outer margin with a row of 8–10 spine-like setae; length of setae ~ 0.2 × maximum width of femur. Apex rounded, with pair of spine-like setae and short, stout, apically blunt setae. Stout, lanceolate, pointed setae scattered along inner margin; femoral patch well developed on fore and middle leg, reduced or rudimentary on hind leg. Tibia. Outer margin with row of short, stout, apically rounded setae, distalmost seta larger. Inner margin with row of medium spine-like setae; on apex tuft of fine, simple setae. Patella-tibial suture present on basal 1 / 2. Tarsus. Outer margin with row of short, stout, apically rounded setae. Inner margin with row of curved, spine-like setae increasing in length distally. Claw with one row of 12 or 13 denticles; distally pointed. Abdominal terga (Fig. 6 f, g). Surface with irregular rows of U-shaped scale bases and fine, simple, scattered setae. Posterior margin of terga: I – IX with triangular spines, wide and short on I, less wide and longer toward end of abdomen; terga VIII and IX additionally with posterolateral spines. Abdominal sterna. Posterior margin of sterna: I – VI smooth, without spines; VII – IX with small, triangular spines. Tergalii (Fig. 6 j, k). Present on segments I – VII. Margin with small denticles intercalating fine, simple setae. Tracheae limited to trunk and few branches. Tergalius I as long as length of 1 / 2 II; tergalius IV as long as segments V and 1 / 3 VI combined; tergalius VII approx. as long as segment VIII. Paraproct (Fig. 6 h). Distally not expanded, with ~ 14 stout, marginal spines. Surface scattered with U-shaped scale bases and fine, simple setae. Cercotractor with numerous small, marginal spines. Imago. Unknown. Etymology. The species is dedicated to the indigenous Lahu people in Thailand. Distribution. Thailand (Fig. 32 a).
Papuanatula (Papuanatula) batantaraja Kovács, Kaltenbach & Gattolliat sp. nov. Figs 6, 7, 8 Type material. Holotype. Indonesia • larva; West Papua, Batanta Island, right side stream of Kalijakut River; 00°52'16"S, 130°37'45.5"E; 545 m; 10. ii. 2024; leg. T. Kovács and R. Sauyai; on slides; GBIFCH 00975887, GBIFCH 01221819; 2024-13.b, EPHTYP-1; MM. Paratypes. 4 larvae; same data as holotype; 2 on slides; GBIFCH 00975888, GBIFCH 01221825, GBIFCH 01221826; 2 in alcohol; GBIFCH 00975886; MZL • 13 larvae; partly same data as holotype; 21. ii. 2025; leg. T. Kovács; 2 in alcohol; GBIFCH 01582004; MZL; 11 in alcohol; 2025-19, EPHTYP-2; MM • 1 larva; West Papua, Batanta Island, right side stream of Kalijakut River, side spring area; 00°52'27"S, 130°37'51"E; 432 m; 20. ii. 2025; leg. T. Kovács; in alcohol; 2025-18.b, EPHTYP-3; MM. Diagnosis. Larva. The following combination of characters distinguishes P. batantaraja sp. nov. from other species of Papuanatula s. str.: large species (> 7 mm body length); body dorsally without row of long, fine, simple setae along midline; fore protoptera posteromedially with pair of minute, broad protuberances and one minute, rather pointed protuberance between them; metanotum and abdominal terga I – X medially with conspicuous, long, pointed protuberance, slightly bent posteriad; left mandible without subtriangular process; femur anteriorly with angulate blank in basal ½; transparent, semicircular scales on anterior surface of femur and dorsal surface of abdominal terga; paracercus vestigial. Description. Larva (Figs 6 – 8). Body length 7.0– 7.5 mm, largest Papuanatula species known so far. Cerci ~ 2 × as long as body length. Cuticular coloration (Figs 6 a – c, 8 e). Head, thorax and abdomen dorsally uniform grey-brown. Head, thorax and abdomen ventrally beige. Legs yellow-brown, femur basally with angulate blank, apically blank. Caudalii yellow-brown. Hypodermal coloration (Fig. 6 a, b). Thorax dorsally with few blackish markings. Abdominal terga with narrow dark brown to blackish transverse band along posterior margins; sigillae blackish. Head. Antenna (Fig. 7 j). Length ~ 1.5 × head length. Flagellum distally with brown dots. Developing turbinate eyes in last instar male larva unknown. Labrum (Fig. 7 a, b). Very wide, length 0.5 × maximum width, laterally angulate. Dorsal, sub-marginal arc with> 40 densely articulated, feathered setae. Right mandible (Fig. 7 d). Margin between prostheca and mola straight, smooth. Otherwise, as typical for the subgenus. Left mandible (Fig. 7 c). Margin between prostheca and mola straight, smooth; subtriangular process not developed. Otherwise, as typical for the subgenus. Hypopharynx (Fig. 7 f). Apical tuft of spine-like setae laterally denser, giving the impression of a pair of tufts. Otherwise, as typical for the genus. Maxilla (Fig. 7 g). Maxillary palp slightly longer than galea-lacinia; palp segment II approx. as long as segment I; segment I thicker than segment II. Otherwise, as typical for the genus. Labium (Fig. 7 e). Paraglossa dorsally with one spine-like seta near inner, distolateral margin. Labial palp with segment I 0.9 × length of segments II and III combined. Segment II with very small, rounded, distomedial protuberance, dorsally with row of five spine-like setae near outer, distolateral margin. Segment III slightly pentagonal, pointed, 0.8 × length of segment II. Otherwise, as typical for the genus. Thorax. Sterna. Protuberances not developed. Terga (Fig. 8 e). Metanotum posteromedially with long, pointed protuberance, slightly bent posteriad. Fore protoptera posteromedially with pair of minute, broad protuberances and one minute, rather pointed protuberance between them. Legs (Fig. 8 a – d). Ratio of leg segments: fore leg 1.1: 1.0: 0.3: 0.1, middle leg 1.0: 1.0: 0.3: 0.1 and hind leg 1.1: 1.0: 0.3: 0.1. Femur. Length ~ 3 × maximum width. Anterior surface with transparent, semicircular scales. Claw with one row of 7–9 denticles, apical denticle larger than other ones, and one or two posterior setae, and one or two short, reduced setae opposite to posterior setae. Otherwise, as typical for the subgenus. Abdomen. Terga (Figs 6 a, b, 8 e – g). Abdominal terga I – X posteromedially with conspicuous, long, pointed protuberance, slightly bent posteriad; Posterior margin of terga: I – IX with minute, pointed denticles, slightly increasing in size toward IX. Surface with scattered transparent, semicircular scales. Tergalii (Fig. 7 h, i). Present on terga II – VII. Broad oblique ovoid; tracheation strongly developed; with grey pigmentation, especially in anal ½; margins smooth, with many short, fine, simple setae. Tergalius II as long as abdominal terga III and IV combined, tergalius IV as long as terga V, VI and ⅓ VII combined, tergalius VII reaching beginning of tergum X. Paraproct (Fig. 8 j). Posterior margin expanded, smooth. Caudalii (Fig. 8 h, i) Cerci without swimming setae. Paracercus vestigial. Pose of subimaginal gonostyli under larval cuticle. Unknown. Subimago. Unknown. Imago. Unknown. Egg. Unknown. Biological aspects. The species is known from the highest watercourse of Batanta, the Kalijakut River system, on altitudes between 430 m and 545 m. The upper habitat (Fig. 29 a, type locality) has volcanic bedrock, the side branch is very fast-flowing, the larvae live on the stones of the steep run. It co-occurs with larvae of P. batanlenos sp. nov. and P. cukiclara sp. nov. In the lower habitat (Fig. 29 b), the species is living in fast flowing water on steep, calcareous surfaces, associated with P. cukiclara sp. nov. Etymology. The species name batantaraja refers to the Indonesian island Batanta, where the species was found, and the Indonesian word “ raja ” meaning king, because it is the largest species known in the genus Papuanatula. Distribution. New Guinea, Batanta Island (Fig. 30).
Papuanatula (Papuanatula) arfak Kaltenbach, Kovács & Gattolliat sp. nov. Figs 1, 2, 3, 4, 5 Type material. Holotype. Indonesia • larva; New Guinea, Papua Prov., Riv. Je, Loc. Arfak, E of Amber village; 01°06'35"S, 133°56'51"E; 1200 m; 16. vi. 2016; leg. B. Sumoked and M. Balke; (BH 68); on slide; GBIFCH 00592543; MZB. Paratypes. 24 larvae; same data as holotype; 4 on slides; GBIFCH 00975788, GBIFCH 00975789, GBIFCH 00592534, GBIFCH 00592540, GBIFCH 01221764; MZL; 20 in alcohol; GBIFCH 00975790, GBIFCH 00976060, GBIFCH 00976063, GBIFCH 00976111; MZL. Diagnosis. Larva. The following combination of characters distinguishes P. arfak sp. nov. from other species of Papuanatula s. str.: body dorsally with irregular row of long, fine, simple setae along midline; abdominal terga without distinct protuberances; abdominal terga I – VI with hypodermal wide dark brown transverse band close to anterior margin, I – IX with medial narrow, dark brown, longitudinal streak; femur proximally with wedge-shaped blank; abdominal terga with triangular, apically rounded denticles on posterior margin; small scattered scales on abdominal terga oblong, striated, apically denticulate; paracercus with nine segments. Description. Larva (Figs 1 – 5). Body length 4.5–5.7 mm, cerci much longer than body length (~ 1.4 ×). Cuticular coloration (Figs 1 a – c, 3 a). Head, thorax and abdomen dorsally brown; thorax with indistinct, complex pattern. Femur proximally with wedge-shaped blank, surrounded by grey area, distally yellow-brown to grey-brown; tibia grey; tarsus grey-brown, distally darker. Head, thorax and abdominal segment I ventrally ecru, protuberances of thoracic sterna brown; abdominal segments II – X ventrally brown, laterally paler. Cerci yellow-brown. Hypodermal coloration (Fig. 1 a). Abdominal terga I – VI (VII) with wide dark brown transverse band close to anterior margin, I – IX with narrow dark brown transverse band close to posterior margin and medially narrow dark brown, longitudinal streak. Head (Figs 1 b, 3 h). Dorsally with irregular row of long, fine, simple setae along midline. Antenna. Length ~ 1.5 × head length. As typical for subgenus. Developing turbinate eyes in last instar male larva (Fig. 3 h) large, subquadrangular, touching each other in the middle. Labrum (Fig. 2 a, b). Length 0.5 × maximum width, laterally convex. Dorsal, sub-marginal arc with ~ 30 feathered setae. Right mandible (Fig. 2 c). Margin between prostheca and mola straight, with row of minute denticles. Otherwise, as typical for subgenus. Left mandible (Fig. 2 d). Margin between prostheca and mola straight, with row of minute denticles. Otherwise, as typical for subgenus. Hypopharynx (Fig. 2 f). As typical for genus. Maxilla (Fig. 2 g, h). Maxillary palp subequal in length to galea-lacinia; palp segment II slightly longer than segment I. Otherwise, as typical for the genus. Labium (Fig. 2 e). Paraglossa dorsally with two spine-like setae near inner, distolateral margin. Labial palp with segment I subequal in length to segments II and III combined. Segment II with minute distomedial protuberance, dorsally with row of five or six spine-like setae near outer, distolateral margin. Segment III slightly pentagonal, pointed; 0.8 × length of segment II. Otherwise, as typical for the genus. Thorax. Sterna (Fig. 1 c). With small protuberances on sides of prosternum and close to openings of mesothoracic and metathoracic sternal apodemes. Terga (Fig. 1 b) without protuberance; with irregular row of long, fine, simple setae along midline. Legs (Fig. 3 a – g). Ratio of leg segments: fore leg 0.9: 1.0: 0.3: 0.2, middle leg 1.0: 1.0: 0.3: 0.2 and hind leg 1.1: 1.0: 0.3: 0.2. Femur. Length ~ 4 × maximum width. Claw with one row of 5–8 denticles and one posterior setae. Otherwise, as typical for subgenus. Abdomen. Terga (Figs 1 b, 4 a, 5 a, b) with irregular row of long, fine, simple setae along midline. Terga without protuberances, terga I – IV with slight, paired medioposterior elevations. Posterior margin of terga: I smooth, without denticles, II – IX with triangular, partly apically rounded denticles, becoming longer toward end of abdomen. Surface with scattered small, oblong, striated, apically serrate scales. Tergalii (Fig. 4 e, f). Present on terga II – VII. Broad ovoid, tracheation well developed and pigmented; margin smooth, with short, fine, simple setae. Tergalius II nearly reaching end of tergum IV, tergalius IV as long as length of terga V and VI combined, tergalius VII nearly reaching end of tergum IX. Paraproct (Fig. 4 g – i). Posterior margin with prolongation and dense row of minute denticles. Caudalii (Fig. 4 b – d). Cerci apart from basal part with 1–6 swimming setae per segment, increasing toward distal part. Paracercus with nine segments. Pose of subimaginal gonostyli under larval cuticle (Fig. 2 i) as typical for the subgenus. Subimago. Unknown. Imago. Unknown. Egg. Unknown. Biological aspects. The species was found at an altitude of 1200 m, together with Papuanatula dumspinae Kaltenbach, Kluge & Gattolliat, 2025, Papuanatula paratuber Kaltenbach, Kluge & Gattolliat, 2025, Papuanatula epituber Kaltenbach, Kluge & Gattolliat, 2025, and Papuanatula pilosa Kaltenbach, Kluge & Gattolliat, 2025. Etymology. The species name refers to the locality Arfak, the type locality of the species. Distribution. New Guinea (Fig. 30).
