Nymphs of the genus Thraulus are adequately characterised by Grant (2024): two dorsal rows of setae on the dorsal face of labrum; width of labrum subequal to width of clypeus (range: 0.81–1.07); lateral margins of clypeus parallel; posterolateral projections on abdominal segments VI–IX, VII–IX or VIII–IX; gills I–VII dissimilar; gill I composed of one or two lanceolate and long lamellae, or a dorsal lanceolate lamella and a ventral oval and fringed lamella; gills II–VII composed of two fringed lamellae along the entire margin, with some species having the basal half of lateral margin bare. Contrary to Grant (2024) the ventral side of labrum can bear not only a row of short stout setae (e.g., Fig. 12B) but in some species scattered stout or thin setae (e.g., Fig. 18B). The hypopharynx does not always bear a “small, rounded, posterolateral projection on arms of the superlingua”, compare for instance Fig. 21F and Fig. 29F.

The species Thraulus eatoni Grant, 2024 [Indonesia Sulawesi], Thraulus parentalis Grant, 2024 [peninsular Malaysia], Thraulus petersorum Grant, 2024 [peninsular Malaysia, east Malaysia Sabah], Thraulus semicastaneus (Gillies, 1951) [southern India], Thraulus thraker Jacob, 1988 [Bulgaria] cannot be assigned to a subgenus because their nymphs are still unknown.

Identical to the generic diagnosis, except that gill I is always composed of one or two lanceolate and long lamellae.

Thraulus bellus Eaton, 1881 by original designation.

• Thraulus (Thraulus) amravati Vasanth, Subramanian & Selvakumar, 2022 [southern India],
• T. (T.) bellus Eaton, 1881 [Europe],
• T. (T.) bishopi Peters & Tsui, 1972 [Peninsular Malaysia, Vietnam, southeastern China],
• T. (T.) connubialis Grant, 2024 [east Malaysia Sabah],
• T. (T.) cursus Grant, 2024 [Japan],
• T. (T.) cuspidatus Vasanth, Subramanian & Selvakumar, 2022 [southern India],
• T. (T.) demoulini Peters & Tsui, 1973 [Thailand],
• Th. (Th.) eloisae sp. nov. [Papua New Guinea],
• T. (T.) fatuus Kang & Yang, 1994 [Taiwan, Japan],
• T. (T.) femoratus Li, Liu & Zhou, 2006 [southeastern China],
• Th. (Th.) granti sp. nov. [Papua New Guinea],
• T. (T.) ishiwatai Grant, 2024 [Japan Okinawa],
• T. (T.) jacobusi Srinivasan, Sivaruban & Barathy, 2022 [southern India],
• Th. (Th.) longinquus sp. nov. [Papua New Guinea],
• T. (T.) macilentus Kang & Yang, 1994 [Taiwan, Japan],
• T. (T.) madagasikarensis Grant, 2024 [Madagascar],
• T. (T.) mudumalaiensis Soman, 1991 [southern India],
• Th. (Th.) nabire sp. nov. [Indonesia, Papua],
• T. (T.) nihonensis Grant, 2024 [Japan Okinawa],
• Th. (Th.) noe sp. nov. [Papua New Guinea],
• Th. (Th.) ogea sp. nov. [Papua New Guinea],
• T. (T.) plumeus Selvakumar, Vasanth & Subramanian, 2022 [northern India],
• Th. (Th.) timorensis sp. nov. [Indonesia, West Timor],
• T. (T.) turbinatus (Ulmer, 1909) [Comoros],
• T. (T.) umbrosus Kang & Yang, 1994 [Taiwan],
• T. (T.) vellimalaiensis Vasanth, Subramanian & Selvakumar, 2022 [southern India],
• Th. (Th.) wemale sp. nov. [Indonesia, Seram].

Mainly in the Oriental and Palaearctic regions, with only two species known from the Afrotropical area in the islands of Comoros and Madagascar; the subgenus seems absent from continental Africa south of the Sahara, as well as from Australia.

Holotype. • Papua New Guinea, nymph on slide, Eastern Highlands Province, Marawaka, Ande, 1700–1800 m, 09.XI.2006, 07°01.697'S, 145°49.807'E, M. Balke and Kinibel col [PNG87], GBIFCH01223087 (ZSM). Paratypes. 3 nymphs in ethanol, GBIFCH01523471, 2 nymphs entirely mounted on slide, GBIFCH01223090, GBIFCH01223089, same data as holotype (MZL). • Papua New Guinea, 1 nymph mounted on slide, GBIFCH01223088, Central Province, Woitape, 1500 m, I.2008, 08°33.178'S, 147°15.481'E, Posman col [PNG167] (MZL).

Body length, male: 6–6.5 mm; female: 7–7.5 mm.

Labrum rectangular, with emargination shallow; proximo-lateral margin of stipes with a single long and stout seta; maxillary palp greatly elongated; outer margin of maxillary palp segment II without stout setae; inner margin of labial palp segment I with 16–18 stout setae; apex of superlingua truncate. Gills II–VI completely fringed with filaments, with ~ 30 filaments on dorsal lamella and ~ 40 on ventral lamella; posterior margin of abdominal segment VIII without denticles; posterolateral projections of the abdomen present on segments VIII–IX.

Head yellowish, greyish between ocelli. Antenna yellowish. Compound eye lower portion black, upper portion brownish orange. Mouth parts pale brown. Thorax. Pronotum pale brown, with greyish areas laterally and medially as in Fig. 2A. Mesonotum greyish brown with scattered lighter maculae. Femur fore femur greyish, mid- and hind femur whitish, distal 1/3 washed with grey. Tibia yellowish orange, with apical macula greyish. Tarsi orange to yellow. Abdomen. Terga evenly greyish brown, lighter along the sagittal line. Sterna pale brown. Gills pale brown to greyish brown. Terminal filaments uniformly pale brown.

Head. Labrum rectangular (Fig. 3A). Ratio labrum width/clypeus width 0.87–0.90. Ratio labrum width/insertion width 1.08–1.10. Medial emargination shallow with rounded denticles (Fig. 3B). Proximal row of setae present, simple. Number of setae on proximal row ~ 75–100. Distal row of setae present, simple. Median part of labrum in ventral view with stout setae in row (Fig. 3B). Mandibles. Outer margin with row of setae on distal 2/3. Ending of the row before incisor. Right mandible with 10 or 11 setae below the mola (Fig. 3C). Maxillae. Posterior margin of cardo with few long hair-like setae, together with some long and stout setae (Fig. 34A). Proximo-lateral margin of stipes with a single long and stout seta. Apical-ventral row with 15–18 pectinate setae. Maxillary palp 1.5 × longer than maxilla; maxillary palp segment II 1.17–1.39 × as long as segment I. Maxillary palp segment III 0.77–0.94 as long as segment I (Fig. 3D). Few stout setae on outer margin of segment I. Segment II inner margin with one stout seta. Segment II outer margin without stout setae. Segment III inner margin with three or four stout setae, dorsal surface fully covered with setae, 2.57–2.64 × as long as base width. Labium. Labial palp segment II 0.59–0.71 as long as segment I. Labial palp segment III 0.64–0.71 as long as segment I (Fig. 3E). Segment I inner margin with 16–18 stout setae, outer margin with 12–15 stout setae. Segment II outer margin with two or three stout setae. Segment III dorsal face with six or seven stout setae (Fig. 3F), 2.00–2.43 × as long as base width. Hypopharynx apex of superlingua truncate.

Thorax. Dorsal margin of femur with a row of long, stout, pointed setae; submarginal row with setae as long as those of dorsal margin (Fig. 4A); ventral margin with short, blunt, stout setae and a submarginal row of long, pointed, stout setae, except on hind leg where they are short and blunt (Fig. 4B); central area of upper surface with few long pointed setae. Fore tibia with several rows of numerous simple feathered setae on ventral margin; outer margin with few hair-like setae; middle tibia with two rows of long pointed setae on ventral margin, outer margin with few hair-like setae; hind tibia with a row of long thin setae on ventral margin, upper and lower surfaces covered with simple and some feathered setae, outer margin with short and long, pointed, stout setae. Tarsal claw straight with 5–8 minutes denticles subequal in length (Fig. 4C).

Abdomen. Gill I with single bifid lamella. Gills II to VII with dorsal and ventral lamellae, ventral lamella smaller. Gills II–VI with upper lamella lacking filaments on basal half of outer margin, with ~ 25–30 short filaments, lower lamella with ~ 35–40 short filaments (Fig. 4E); on both lamellae apical filament much longer than the others; gill VII small, with narrow lamellae ending with a single filament (Fig. 4F). Posterior margin of terga IX with denticles (Fig. 4D). Posterior margin of terga V–VIII without denticles. Posterolateral projections present on segments VIII and IX.

This species is dedicated to Eloïse Sartori, daughter of the first author.

Holotype. • Papua New Guinea, nymph on slide, Enga Province, Kumul Lodge at foot of Mt Hagen, 2700 m, 05.XII.2006, 05°47.548'S, 143°58.761'E, M. Balke and Kinibel col [PNG124], GBIFCH01223092 (ZSM). Paratypes. 4 nymphs in ethanol, GBIFCH01523472, 1 nymph entirely mounted on slide, GBIFCH01223093, same data as holotype (MZL). • Papua New Guinea, one nymph on slide, GBIFCH01223094, Central Province, Kokoda trek, 1390 m, I.2006, 09°00.338'S, 147°44.252'E, Posman col. [PNG173] (MZL).

Body length, female: 10.5–11 mm.

Labrum cordiform, medial emargination shallow with flat denticles; dorsal face of labrum with a simple row in distal position; median part of labrum in ventral view with stout setae in row; apical ventral row of maxilla composed of 19–23 pectinate setae; tarsal claw with 7–10 teeth; gill lamellae on segments II–VI with > 25 filaments each; posterolateral projections of the abdomen present on segments VII–IX.

Head. Pale brown, medium brown between ocelli. Antenna broken. Mouth parts pale brown. Thorax. Pronotum yellowish brown, washed with greyish maculae laterally and medially as in Fig. 2B. Mesonotum greyish brown with scattered lighter maculae; anterolateral base of forewing pads with a blackish dot. Femur yellowish with distal 1/3 greyish; forefemur also with a middle greyish band. Tibia yellowish, apex of fore tibia greyish. Tarsi yellowish. Abdomen. Terga evenly greyish brown with a pair of lighter maculae on segments II–VII; segments VIII and IX darker, segment X lighter. Sterna yellowish grey in the middle, yellowish on sides, segments VIII and IX darker, brownish. Gills greyish brown, upper lamella darker. Terminal filaments uniformly yellowish.

Head. Labrum cordiform (Fig. 5A). Ratio labrum width/clypeus width 0.90–0.95. Ratio labrum width/insertion width 1.23–1.24. Medial emargination shallow with flat denticles. Proximal row of setae present, simple. Number of setae on proximal row ~ 26–40. Distal row of setae present, simple. Median part of labrum in ventral view with stout setae in row (Fig. 5B). Mandibles. Outer margin with row of setae on distal 2/3. Ending of the row near incisor. Right mandible with 10–17 setae below the mola (Fig. 5C). Maxillae. Posterior margin of cardo with few hair-like setae and three or four stout and long setae in submarginal position. Proximo-lateral margin of stipes with a single long and stout seta. Apical-ventral row with 19–23 pectinate setae. Maxillary palp segment II 0.89–1.00 as long as segment I. Maxillary palp segment III 0.67–0.80 as long as segment I (Fig. 5D). Stout setae on outer margin of segment I absent. Segment II inner margin with 3–7 stout setae, outer margin with 3–5 stout setae. Segment III inner margin with four or five stout setae, dorsal surface with few setae at apex, 1.60–2.10 × as long as base width. Labium. Labial palp segment II 0.69–0.74 as long as segment I. Labial palp segment III 0.96–0.86 as long as segment I (Fig. 5E). Segment I inner margin with 9–15 stout setae, outer margin with 15–20 stout setae. Segment II outer margin with two to four stout setae. Segment III dorsal face with five or six setae, 3.00–3.38 × as long as base width. Hypopharynx. Apex of superlingua truncate (Fig. 5F).

Thorax. Dorsal margin of femur with a row of long, stout, pointed setae; irregular submarginal row with setae as long as or half the size of those of dorsal margin (Fig. 6A); ventral margin with shorter, stout setae and a submarginal row of long and pointed stout setae, except on hind leg where those on ventral margin are short and blunt (Fig. 6B); central area of upper surface with few long, pointed setae somewhat arranged in a longitudinal row. Fore tibia with several rows of numerous simple, feathered setae on ventral margin; outer margin of fore tibia with few hair-like setae; middle tibia with two rows of long, pointed setae on ventral margin, outer margin with numerous hair-like setae; hind tibia with a row of long thin setae on ventral margin, upper and lower surface covered with simple and some feathered setae, outer margin with short and long, pointed, stout setae. Tarsal claw straight with 7–10 denticles progressively larger apically (Fig. 6C).

Abdomen. Gill I with single bifid lamella. Gills II–VII with dorsal and ventral lamellae, dorsal lamella smaller. Gill II–VI with upper and lower lamella entirely bordered with ~ 30 short filaments (Fig. 6F); gill VII with ~ 15 filaments and ~ 10 filaments on upper and lower lamellae, respectively (Fig. 6G). Posterior margin of terga IX with minute denticles (Fig. 6D), or with small denticles. Posterior margin of terga VIII with minute denticles (Fig. 6E). Posterior margin of terga V–VII without denticles. Posterolateral projections present on segments VII–IX.

This species is dedicated to Prof. Peter Grant (SWOSU, Oklahoma, USA) for his outstanding contribution to the study of the genus Thraulus.

Holotype. • Papua New Guinea, nymph on slide, Madang Province, Aiome area, 130 m, 11.III.2007, 05°10.593'S, 144°42.800'E, Kinibel col [PNG156], GBIFCH01223098 (ZSM). Paratype. • Papua New Guinea, one nymph in ethanol, GBIFCH01523473, same data as holotype (MZL).

Body length, male: 5.5 mm.

Labrum cordiform, medial emargination shallow with flat denticles; dorsal face of labrum with a simple row in distal position; median part of labrum in ventral view with stout setae in row; apical ventral row of maxilla composed of 17 pectinate setae; tarsal claw with six teeth; gill lamellae on segments II–VI with > 25 filaments each; posterolateral projections of the abdomen present on segments VII–IX.

Head. Yellowish brown, medium brown between ocelli. Antenna yellowish, pedicel dark brown. Compound eye lower portion black, upper portion medium brown. Mouth parts yellowish brown. Thorax. Pronotum pale brown washed with medium brown. Mesonotum pale brown washed with medium brown. Femur whitish, apex tinted with dark brown. Tibia whitish. Tarsi yellowish. Abdomen. Terga uniformly dark brown, sagittal line somewhat paler. Sterna greyish brown. Gills lamellae pale grey, filaments greyish purple. Terminal filaments broken.

Head. Labrum cordiform (Fig. 7A). Ratio labrum width/clypeus width 0.91. Ratio labrum width/insertion width 1.21. Medial emargination shallow with flat denticles. Proximal row of setae present, simple. Number of setae on proximal row ~ 24. Distal row of setae present, simple. Median part of labrum in ventral view with stout setae in row (Fig. 7B). Mandibles. Outer margin with middle and distal tuft of setae. Right mandible with 12 setae below the mola (Fig. 7C). Maxillae. Posterior margin of cardo broken. Proximo-lateral margin of stipes with a single long and stout seta. Apical-ventral row with 17 pectinate setae. Maxillary palp segment II 0.82 as long as segment I. Maxillary palp segment III 0.75 as long as segment I (Fig. 7D). Stout setae on outer margin of segment I absent. Segment II inner margin with three stout setae. Segment II outer margin without stout setae. Segment III inner margin with four stout setae. Segment III dorsal surface with few setae at apex. Segment III 1.60 × as long as base width. Labium. Labial palp segment II 0.68 as long as segment I. Labial palp segment III 0.76 as long as segment I (Fig. 7E). Segment I inner margin with nine stout setae, outer margin with eight stout setae. Segment II outer margin with four stout setae. Segment III dorsal face with four stout setae. Segment III 3.11 × as long as base width. Hypopharynx. Apex of superlingua pointed (Fig. 7F).

Thorax. Dorsal margin of femur with a row of long, stout, pointed setae; irregular submarginal row of shorter pointed setae; ventral margin with short stout setae and a submarginal row of long, pointed, stout setae, except on hind leg where those on ventral margin are very short and blunt (Fig. 8A); central area of upper surface with minute pointed setae. Middle tibia with two rows of long, simple, feathered, or pointed setae on ventral margin, outer margin with few hair-like setae; hind tibia with row of simple long and thin setae on ventral margin, upper and lower surface covered with simple and feathered setae, outer margin with short and long pointed, stout setae and hair-like setae. Tarsal claw straight with six denticles progressively larger apically (Fig. 8B).

Abdomen. Gill I with single bifid lamella. Gills II–VII with dorsal and ventral lamellae of the same size. Gill II–VI with upper lamella lacking filaments on basal half of outer margin, with ~ 25–30 filaments on each lamella (Fig. 8C); gill VII with ~ 15 filaments on each lamella. Posterior margin of terga IX with denticles. Posterior margin of terga VIII with minute denticles. Posterior margin of terga V–VII without denticles. Posterolateral projections present on segments VII–IX.

The Latin adjective longinquus means distant or lost, as an indication of the remote place where the species was found.

Holotype. • Indonesia, Papua, nymph on slide, Road Nabire-Enarotali KM 62, 340 m, 22.X.2011, 03°31.684'S, 135°42.802'E, M. Balke col [PAP11], GBIFCH01223096 (MZB). Paratypes. • Indonesia, Papua, 1 nymph in ethanol, GBIFCH01523574, 1 nymph mounted on slide, GBIFCH01223097, same data as holotype (MZL).

Body length, male: 4.5 mm.

Labrum cordiform, medial emargination shallow with flat denticles; median part of labrum in ventral view with stout setae in a row; proximal row of setae on dorsal face of labrum with 19–23 setae; apical ventral row of maxilla composed of 15–17 pectinate setae; tarsal claw with eight or nine teeth, distal one much larger; gill lamellae on segments II–VI fringed only on distal half, with < 20 filaments on each.

Head. Greyish brown, clypeus medium brown. Antenna yellowish. Compound eye lower portion black, upper portion brownish red. Mouth parts pale brown, labrum and mandibles medium brown. Thorax. Pronotum yellowish brown, greyish brown on edges and in the middle. Mesonotum pale brown, washed with greyish maculae. Femur whitish with distal 1/3 greyish. Tibia yellowish. Tarsi yellowish. Abdomen. Terga: segments II–VIII medium brown, posterior 1/3 dark brown, segment IX–X medium brown. Sterna greyish brown, posterior 1/3 medium brown. Gills greyish brown to greyish purple. Terminal filaments broken.

Head. Labrum cordiform (Fig. 9A). Ratio labrum width/clypeus width 0.81–0.90. Ratio labrum width/insertion width 1.15–1.26. Medial emargination shallow with flat denticles. Proximal row of setae present simple. Number of setae on proximal row ~ 19–23. Distal row of setae present, simple. Median part of labrum in ventral view with stout setae in row (Fig. 9B). Mandibles. Outer margin with middle and distal tuft of setae. Right mandible with eight to nine setae below the mola (Fig. 9C). Maxillae. Posterior margin of cardo with few hair-like setae and one or two stout and long setae in submarginal position. Proximo-lateral margin of stipes with a single long and stout seta. Apical-ventral row with 15–17 pectinate setae. Maxillary palp segment II 0.88–0.90 as long as segment I. Maxillary palp segment III 0.76–0.85 as long as segment I (Fig. 9D). Stout setae on outer margin of segment I absent. Segment II inner margin with three or four stout setae. Segment II outer margin with 3–5 stout setae. Segment III inner margin with 2–4 stout setae. Segment III 1.86–2.00 × as long as base width. Labium. Labial palp segment II 0.78–0.81 as long as segment I. Labial palp segment III 0.69–0.74 as long as segment I (Fig. 9E). Segment I inner margin with 8–13 stout setae, outer margin with nine stout setae. Segment II outer margin with three or four stout setae. Segment III dorsal face with four or five stout setae. Segment III 2.86–3.29 × as long as base width. Hypopharynx. Apex of superlingua somewhat pointed (Fig. 9F).

Thorax. Dorsal margin of femur with a row of long, stout, pointed setae; submarginal row with long setae decreasing in size towards apex (Fig. 10A); ventral margin with short and stout setae and a submarginal row of long and pointed stout setae, except on hind leg where those on ventral margin are short and blunt (Fig. 10B); central area of upper surface with few long and pointed setae. Fore tibia with several rows of numerous mostly feathered setae on ventral margin; outer margin of fore tibia with few hair-like setae; middle tibia with two rows of long and simple or feathered, pointed setae on ventral margin, outer margin with numerous hair-like setae; hind tibia with a row of simple short thin setae on ventral margin, upper and lower surfaces covered with simple and feathered setae, outer margin with short and long, pointed, stout setae. Tarsal claw hooked with eight or nine denticles progressively larger, except distal one much larger (Fig. 10C).

Abdomen. Gill I with single bifid lamella. Gills II–VII with dorsal and ventral lamellae, dorsal lamella smaller. Gill II–VI with upper lamella lacking filaments on basal half of outer margin, with < 20 short filaments, lower lamella with < 20 short filaments (Fig. 10D). Posterior margin of terga IX with denticles. Posterior margin of terga VIII without denticles, or with minute denticles. Posterior margin of terga V–VII without denticles. Posterolateral projections present on segments VI–IX.

This species is named after the district of Nabire where it can be found and is a noun in apposition.

Holotype. • Papua New Guinea, nymph on slide, Central Province, Woitape, 700 m, I.2008, 8°31.290'S, 147°13.684'E, Posman col [PNG166], GBIFCH01223114 (ZSM). Paratypes. • Papua New Guinea, 2 nymphs in ethanol (UFVB), 8 nymphs in ethanol, GBIFCH01523475, 3 entirely mounted on slide, GBIFCH01223113, GBIFCH01223112, GBIFCH01223111, same data as holotype (MZL). Other material. • Papua New Guinea, one nymph on slide, GBIFCH01223109, Central Province, Kokoda Trek, 1390 m, I. 2008, 09°00.338'S, 147°44.252'E. Posman col [PNG173] (MZL). • Papua New Guinea, one nymph on slide, GBIFCH01223110, Eastern Highlands Province, Marawaka, Ande, 1700–1800 m, 9.XI.2006, approx. 07°01.697'S, 145°49.807'E, M. Balke & Kinibel col [PNG87] (MZL).

Body length, male: 5.1–5.2 mm Body length, female: 6.3–6.5 mm.

Labrum rectangular, with emargination narrow; proximo-lateral margin of stipes with one long and stout seta together with two small and stout setae; maxillary palp greatly elongated; outer margin of maxillary palp segment II with 1–3 stout setae; inner margin of labial palp segment I with 20–26 stout setae; apex of superlingua rounded. Gills II–VI without filaments on basal half, with ~ 15 filaments on dorsal lamella and ~ 20 on ventral lamella; posterior margin of abdominal segment VIII with minute denticles.

Head. Vertex greyish brown, darker between ocelli. Antenna yellowish, pedicel greyish. Compound eye lower portion black, upper portion reddish brown. Mouth parts and clypeus medium brown. Thorax. Pronotum greyish brown. Mesonotum pale brown washed with greyish brown laterally (Fig. 11A). Metanotum dark brown. Femur: forefemur almost completely greyish brown; mid- and hind femora whitish, apical 1/3 brownish. Tibia yellowish, apex yellowish brown. Tarsi yellowish brown. Abdomen. Terga dark brown, posterior margin blackish. Sterna yellowish brown in females to dark brown in males. Gills purplish brown. Terminal filaments uniformly yellowish brown.

Head. Labrum rectangular (Fig. 12A). Ratio labrum width/clypeus width 0.88–0.92. Ratio labrum width/insertion width 1.12–1.13. Medial emargination narrow with rounded denticles. Proximal row of setae present, simple. Number of setae on proximal row 69–87. Distal row of setae present, simple. Median part of labrum in ventral view with stout setae in a row (Fig. 12B). Mandibles. Outer margin with row of setae on distal 2/3, row ending before incisor. Right mandible with 7–10 setae below the mola (Fig. 12C). Maxillae. Posterior margin of cardo broken. Proximo-lateral margin of stipes with one long stout seta together with two small stout setae. Apical-ventral row with 15–17 pectinate setae. Maxillary palp extremely long, almost 2 × longer than maxilla; segment II 1.3–1.8 × as long as segment I. Maxillary palp segment III 0.77–0.99 as long as segment 1 (Fig. 12D). Stout setae on outer margin of segment I present. Segment II inner margin with two stout setae, outer margin with 1–3 stout setae. Segment III inner margin with three or four stout setae, dorsal surface with few setae at apex. Segment III 2.46–3.00 × as long as base width. Labium. Labial palp segment II 0.67–0.71 as long as segment 1. Labial palp segment III 0.63–0.70 as long as segment 1 (Fig. 12E). Segment I inner margin with 20–26 stout setae, outer margin with 12–15 stout setae. Segment II outer margin with two or three stout setae. Segment III dorsal face with six stout setae. Segment III 2.15–2.29 × as long as base width. Hypopharynx. Apex of superlingua rounded (Fig. 12F).

Thorax. Dorsal margin of femur with a row of long stout setae, and a submarginal row of same setae; ventral margin with shorter and stouter setae (Fig. 13A, B); upper surface with few long and pointed but stout setae. Tarsal claw straight with 4–7 subequal denticles (Fig. 13C).

Abdomen. Gill I with single bifid lamella (Fig. 13F). Gills II–VII with dorsal and ventral lamellae, dorsal lamella smaller. Gills II–VI with ~ 15 filaments on upper lamella and ~ 20 on the ventral one, filamentous projections lacking on proximal half (Fig. 13G); gill VII with three filaments on each lamella, one much longer than the other two (Fig. 13H). Posterior margin of terga IX with denticles (Fig. 13D). Posterior margin of terga VIII with minute denticles (Fig. 13E). Posterior margin of terga V–VII without denticles. Posterolateral projections present only on segment VIII–IX.

Eggs. General shape elongated. Size 190 μm × 75 μm. Chorionic surface smooth, with longitudinal ridges weakly elevated, interrupted partially in the equatorial area. No attachment structures. Micropyle in equatorial position (Fig. 20A).

This species is dedicated to Noe Sartori, grandson of the first author, and is a noun in apposition.

Holotype. • Papua New Guinea, nymph on slide, Madang Province, Trans Gogol, 30 m, II.2008, 5°18.0915'S, 145°36.4532'E, BRC col [PNG179], GBIFCH01223108 (ZSM). Paratypes. • Papua New Guinea, 4 nymphs in ethanol (UFVB), 22 nymphs in ethanol, GBIFCH01513476, 2 nymphs entirely mounted on slides, GBIFCH01223107, GBIFCH01223106, same data as holotype (MZL).

Body length, male: 3.6–3.9 mm Body length, female: 4.2–4.4 mm.

Labrum cordiform, medial emargination shallow with flat denticles; median part of labrum in ventral view with stout setae in a row; outer margin of mandibles with tuft of setae in medium and distal positions; posterior margin of cardo with few hair-like setae; gill lamellae on segments II–VI without filaments on basal half, with < 20 filaments on each; posterolateral projections of the abdomen present on segments VII–IX.

Head. Pale brown, greyish brown between ocelli and eyes. Antenna whitish. lower portion black, upper portion reddish brown, outer ommatidia dark brown. Mouth parts: clypeus greyish brown. Thorax. Pronotum greyish brown, with two pale brown markings medially. Mesonotum greyish brown, washed with dark brown laterally and medially (Fig. 11B). Metanotum greyish brown. Femur yellowish brown, with one pale brown marking in the middle and one distally. Tibia uniformly yellowish. Tarsi uniformly yellowish. Abdomen. Terga medium brown, posterior margin dark brown. Sterna yellowish grey. Gills purplish brown. Terminal filaments uniformly yellowish white.

Head. Labrum cordiform (Fig. 14A). Ratio labrum width/clypeus width 0.93–0.94. Ratio labrum width/insertion width 1.29–1.30. Medial emargination shallow with flat denticles. Proximal row of setae present, simple. Number of setae on proximal row 35–47. Distal row of setae present, simple. Median part of labrum in ventral view with only scattered stout setae (Fig. 14B). Mandibles. Outer margin with middle tuft of setae, and with distal tuft of setae. Right mandible with nine or ten setae below the mola (Fig. 14C). Maxillae. Posterior margin of cardo with few hair-like setae. Proximo-lateral margin of stipes with a single stout and medium-sized seta. Apical-ventral row with 25–29 pectinate setae. Maxillary palp segment II 0.77–1.05 as long as segment I. Maxillary palp segment III 0.69–0.73 as long as segment I (Fig. 14D). Stout setae on outer margin of segment I absent. Segment II inner margin with five stout setae, outer margin with five or six stout setae. Segment III inner margin with three stout setae, dorsal surface with few setae at apex. Segment III 1.89–2.57 × as long as base width. Labium. Labial palp segment II 0.81–0.82 as long as segment I. Labial palp segment III 0.71–0.81 as long as segment I (Fig. 14E). Segment I inner margin with 8–11 stout setae, outer margin with 12 or 13 stout setae. Segment II outer margin with four or five stout setae. Segment III, dorsal face with three or four stout setae. Segment III 3.0–3.57 as long as base width. Hypopharynx. Apex of superlingua truncate (Fig. 14F).

Thorax. Dorsal margin of femur with a row of stout and long setae, and a submarginal row with same setae; ventral margin with shorter and stouter setae; upper surface with few long and pointed stout setae (Fig. 15A). Tarsal claw hooked with 4–6 denticles progressively larger apically.

Abdomen. Gill I with single bifid lamella. Gills II–VII with dorsal and ventral lamellae, dorsal lamella smaller. Gill II to VI with seven to ten filaments on upper lamella and 12–14 on the ventral one, filamentous projections lacking on proximal half; gill VII with three or four filaments on each lamella. Posterior margin of terga IX with denticles. Posterior margin of terga IX with denticles (Fig. 15B). Posterior margin of terga VIII with minute denticles. Posterior margin of terga V–VII without denticles. Posterolateral projections present on segments VII–IX.

Eggs. General shape elongated. Size 190 μm × 75 μm. Chorionic surface smooth and uplifted compared to attachment structures (Fig. 20B). Attachment structures composed of nine long slender filaments attached to each pole (Fig. 20C); presence of two transverse filaments at the micropyle opening (Fig. 20D). Micropyle in equatorial position.

This species is named after the ethnic group Ogea, who live close to the Gogol River, and is a noun in apposition.

Holotype. • Indonesia, Timor, nymph on slide, GBIFCH01223095, some gills in ethanol, GBIFCH01523478, Naikliu area, restpools in dry forest,130 m, 03.X.2011, 09°58.425'S, 123°41.439'E, M. Balke col [TIM11] (MZB).

Body length, female: 6.5 mm.

Labrum cordiform, medial emargination shallow without denticles; dorsal face of labrum with multiple rows in distal position; tarsal claw hooked with 15 teeth and a palisade of four other teeth in distal position; gill lamellae on segments II–VI with > 25 filaments each; posterolateral projections of the abdomen present on segments VIII and IX.

Femur, tibia, and tarsi whitish.

Head. Labrum cordiform (Fig. 16A). Ratio labrum width/clypeus width 0.95. Ratio labrum width/insertion width 1.39. Medial emargination shallow without denticles. Proximal row of setae present, simple. Number of setae on proximal row ~ 29. Distal row of setae multiple (Fig. 16C). Median part of labrum in ventral view with stout setae in a row (Fig. 16B). Mandibles. Outer margin with middle tuft of setae. Right mandible with ten setae below mola (Fig. 16D). Maxillae. Posterior margin of cardo broken. Proximo-lateral margin of stipes with single long stout seta (Fig. 34E). Apical-ventral row with 22 pectinate setae. Maxillary palp segment II 0.82 as long as segment I. Maxillary palp segment III 0.78 as long as segment I (Fig. 16E). Stout setae on outer margin of segment I absent. Segment II inner margin with five stout setae, outer margin without stout setae. Segment III inner margin with five stout setae, dorsal surface with few setae at apex. Segment III 1.67 × as long as base width. Labium. Labial palp segment II 0.76 as long as segment I. Labial palp segment III 0.76 as long as segment I (Fig. 16F). Segment I inner margin with two stout setae, outer margin with 12 stout setae. Segment II outer margin with ten stout setae. Segment III dorsal face with five stout setae. Segment III 2.90 × as long as base width. Hypopharynx. Apex of superlingua truncate (Fig. 16F).

Thorax. Dorsal margin of femur with a row of long, stout and pointed setae; regular submarginal row with setae as long as those of dorsal margin (Fig. 17A); ventral margin with short and pointed setae and a submarginal row of short and blunt but stout setae, except on hind leg where those on ventral margin are short and blunt (Fig. 17B); central area of upper surface with few long pointed setae. Middle tibia with two rows of long pointed setae on ventral margin, outer margin with few hair-like setae; hind tibia with a row of long thin setae on ventral margin, upper and lower surfaces covered with simple and some feathered setae, outer margin with long and short pointed stout setae. Tarsal claw hooked with 15 denticles, together with a palisade of four small denticles at apex (Fig. 17C).

Abdomen. Gill I with single bifid lamella. Gills II–VII with dorsal and ventral lamellae, dorsal lamella smaller. Gills II–VI with upper lamella entirely bordered with ~ 35 very long filaments, lower lamella with ~ 30 very long filaments (Fig. 17D); gill VII smaller but with the same number of long filaments. Posterior margin of terga IX with denticles. Posterior margin of terga VIII with minute denticles. Posterior margin of terga V–VII without denticles. Posterolateral projections present on segments VIII–IX.

This species is named after the island from where it comes. Although we have a single nymph in hand, we think important to formally named it, since it possesses distinct characters; moreover, it is the first Leptophlebiidae described from this island and only the second mayfly species (Kaltenbach and Gattolliat 2017).

Holotype. • Indonesia, Maluku, Seram, nymph on slide, Kanikeh, 607 m, 7.IV.2012, 3°06.524'S / 129°28.796E, M. Balke col [AMB07], GBIFCH01223099 (MZB). Paratypes. • Indonesia, Maluku, Seram, 9 nymphs in ethanol, GBIFCH01523477, 2 entirely mounted on slide, GBIFCH01223105, GBIFCH01223102, same data as holotype (MZL). • Indonesia, Maluku, Seram, 1 nymph in ethanol (UFVB), 2 entirely mounted on slide, GBIFCH01223104, GBIFCH01223103, Roho-Kanikeh village, waterholes near Roho, 117 m, 7.IV.2012, 3°01.193'S / 129°23.480'E, M. Balke col [AMB06] (MZL). • Indonesia, Maluku, Seram, 1 nymph entirely mounted on slide, GBIFCH01223100, small puddles Kanikeh-Wainsela, 800–1350 m, 9.IV.2012, 3°08.679'S / 129°28.403'E, M. Balke col [AMB10] (MZL). • Indonesia, Maluku, Seram, 1 nymph entirely mounted on slide, GBIFCH01223101, near Huahulo village, 110 m, 13.IV.2012, 2°59.069'S / 129°20.539'E, M. Balke col. [AMB13] (MZL). • Indonesia, Maluku, Seram, 1 nymph entirely mounted on slide, GBIFCH01223149, Gunung Salahutu, stream, shaded at waterfall, 125 m, 3.IV.2012, 3°33.318'S / 128°17.711'E, M. Balke col. [AMB03] (MZL).

Body length, male: 4.8–5 mm, female: 5.7–6 mm.

Labrum cordiform, medial emargination shallow with flat denticles; median part of labrum in ventral view with stout setae in a row; outer margin of mandibles with row of setae on distal 2/3; posterior margin of cardo with many hair-like setae and one stout long seta in submarginal position; gill lamellae on segments II–VI with < 20 filaments, without filaments on basal half ; posterolateral projections of the abdomen present on segments VI–IX.

Head. Pale brown, darker between ocelli in males, greyish brown in females. Antenna whitish. Compound eye lower portion black, upper portion reddish brown. Mouth parts pale brown to greyish brown. Thorax. Pronotum medium brown, darker in anterior part and laterally (Fig. 11C). Mesonotum whitish brown, darker laterally, with greyish marks along the median suture. Metanotum pale brown. Femur greyish white, dorsal surface with two large dark grey marks proximally and distally. Tibia uniformly pale brown. Tarsi uniformly pale brown. Abdomen. Terga uniformly medium to dark brown, tergites IX and X paler in males. Sterna yellowish grey. Gills purplish brown. Terminal filaments uniformly pale brown.

Head. Labrum cordiform (Fig. 18A). Ratio labrum width/clypeus width 0.96–1.03. Ratio labrum width/insertion width 1.31–1.39. Medial emargination shallow with flat denticles. Proximal row of setae present, simple. Number of setae on proximal row 33–45. Distal row of setae present, simple. Median part of labrum in ventral view with only scattered stout setae (Fig. 18B). Mandibles. Outer margin with row of setae on distal 2/3 ending near incisor. Right mandible with 7–11 setae below the mola (Fig. 18C). Maxillae. Posterior margin of cardo with many hair-like setae and one stout long seta in submarginal position. Proximo-lateral margin of stipes with a single stout long seta. Apical-ventral row with 18–23 pectinate setae. Maxillary palp segment II 0.70–1.00 × as long as segment I. Maxillary palp segment III 0.67–0.82 as long as segment 1 (Fig. 18D). Stout setae on outer margin of segment I absent. Segment II inner margin with four or five stout setae, outer margin with 4–6 stout setae. Segment III inner margin with 4–6 stout setae, dorsal surface with few setae at apex. Segment III 1.8–2.22 × as long as base width. Labium. Labial palp segment II 0.72–0.90 as long as segment I. Labial palp segment III 0.67–0.78 as long as segment I (Fig. 18E). Segment I inner margin with 9–13 stout setae, outer margin with 11 or 12 stout setae. Segment II outer margin with 3–5 stout setae. Segment III dorsal face with 3–5 stout setae. Segment III 3.20–3.57 × as long as base width. Hypopharynx. Apex of superlingua truncate (Fig. 18F).

Thorax. Dorsal margin of femur with a row of stout, long setae, and a submarginal row with same setae; ventral margin with shorter and stouter setae; upper surface with few long, pointed stout setae (Fig. 19A, B). Tarsal claw hooked with 4–7 denticles progressively larger apically (Fig. 19C).

Abdomen. Gill I with single bifid lamella (Fig. 19F). Gills II–VII with dorsal and ventral lamellae, dorsal lamella smaller. Gills II–VI with < 10 filaments on each lamella (Fig. 19G); gill VII with two or three filaments on each lamella (Fig. 19H). Posterior margin of terga IX with denticles (Fig. 19D). Posterior margin of terga VII–VIII with minute denticles (Fig. 19E). Posterior margin of terga V–VI without denticles. Posterolateral projections present on segment VI–IX.

Eggs. General shape elongated. Size 190 μm x 75 μm. Chorionic surface smooth and uplifted compared to attachment structures. Attachment structures composed of 12 long and slender filaments attached to each pole and able to inflate (Fig. 20E). At the micropyle opening, presence of two transverse short filaments, easily broken (Fig. 20F). Micropyle in equatorial position.

This species is named to honour the ancient ethnic group Wemale, who still inhabits the inland of Seram where the specimens of this species were collected and is a noun in apposition.

• Papua New Guinea, nymph on slide, Madang Province, Keki, Adalbert Mts, 400 m, 29.XI.2006, 04°43.058'S, 145°24.437'E, Binatang Boys col [PNG119], GBICH01223091 (MZL).

Nymph. Labrum trapezoidal; ventral side of the labrum with only scattered stout setae; labial palp with segment III > 4 × as long as base width. Posterior margin of abdominal segment VI–VII with well-developed denticles.

Femur whitish, with distal 1/3 greyish; forelegs greyish along the dorsal margin. Tibia whitish, apex greyish. Tarsi yellowish.

Head. Labrum trapezoid (Fig. 21A). Ratio labrum width/clypeus width 1.07. Ratio labrum width/insertion width 1.52. Medial emargination shallow with flat denticles. Proximal row of setae simple, shifted to distal margin (Fig. 21B). Number of setae on proximal row ~ 30. Distal row of setae present, simple. Median part of labrum in ventral view with only scattered stout setae. Mandibles. Outer margin with row of setae on distal 2/3. Ending of the row before incisor. Right mandible with eight setae below the mola (Fig. 21C). Maxillae. Posterior margin of cardo with very few hair-like setae and a long and stout seta in submarginal position. Proximo-lateral margin of stipes with a single long and stout seta. Apical-ventral row with 26 pectinate setae. Maxillary palps lost (Fig. 21D). Labium. Labial palp segment II 0.87 as long as segment I. Labial palp segment III 0.89 as long as segment I (Fig. 21E). Segment I inner margin with five stout setae, outer margin with eight stout setae. Segment II outer margin with two stout setae. Segment III dorsal face with eight stout setae. Segment III 4.40 × as long as width base. Hypopharynx. Apex of superlingua truncate (Fig. 21F).

Thorax. Dorsal margin of femur with a row of long, stout, pointed setae (Fig. 22A); irregular submarginal row on fore femur, regular on mid femur (Fig. 22B), absent on hind femur; ventral margin with short, stout setae and a submarginal row of long, pointed stout setae, except on hind leg where those on ventral margin are short and blunt; central area of upper surface with few long, pointed setae. Fore tibia with several rows of numerous mostly feathered setae on ventral margin; outer margin of fore tibia with few hair-like setae; middle tibia with two rows of long, simple or feathered, pointed setae on ventral margin, outer margin with few hair-like setae; hind tibia with a row of simple short, thin setae on ventral margin, upper and lower surface covered with simple and feathered setae, outer margin with short and long pointed stout setae. Tarsal claw straight with eight or nine denticles progressively larger, with a palisade of four small denticles at apex (Fig. 22C).

Abdomen. Gills lost. Posterior margin of terga VI–IX with denticles (Fig. 23A–D). Posterior margin of terga V with minute denticles (Fig. 23E). Posterolateral projections present on segments VII–IX.

Identical to the generic diagnosis, except that gill I is always composed of one dorsal lanceolate lamella and a large oval fimbriate ventral lamella.

Masharikella fasciatus (Kimmins, 1956) by original designation.

Thraulus (Masharikella) fasciatus (Kimmins, 1956), comb. nov. [Uganda, Zimbabwe], T. (M.) gopalani Grant & Sivaramakrishnan, 1985, comb. nov. [southern India], Th. (M.) iteris sp. nov. [Papua New Guinea], Th. (M.) johannisluci sp. nov. [Papua New Guinea], T. (M.) malabarensis Vasanth, Subramanian & Selvakumar, 2022, comb. nov. [southern India], T. (M.) opifer Grant, 2024, comb. nov. [northern Australia], Th. (M.) pascalae sp. nov. [Indonesia, Papua], Th. (M.) samueli sp. nov. [Papua New Guinea], T. (M.) thiagarajani Balasubramanian & Muthukatturaja, 2019, comb. nov. [southern India], and T. (M.) torrentis (Gillies, 1964), comb. nov. [Tanzania, Angola].

The subgenus exhibits a disjunct distribution in Africa, southern India, New Guinea, and northern Australia. Currently, it is surprisingly not recorded from Oriental southeast Asia, suggesting the species found in New Guinea originate from tropical Australia. This may indicate a Gondwanan origin for the subgenus. Its presence in Madagascar is expected but yet not proved.

Holotype. • Papua New Guinea, nymph on slide, Central Province, Kokoda Trek, 980 m, I.2008, 09°15.933'S, 147°36.590'E, Posman col [PNG169], GBIFCH01223115 (ZSM). Paratypes. • Papua New Guinea, 1 nymph entirely mounted on slide, GBIFCH01223116, 1 nymph in ethanol, GBIFCH01523480, same data as holotype (MZL). • Papua New Guinea, 1 nymph entirely mounted on slide, GBIFCH01223117, some gills in ethanol, GBIFCH01523479, Central Province, Kokoda Trek, 320 m, I.2008, 09°19.236'S, 147°31.791'E, Posman col [PNG168] (MZL).

Body length, female: 5.8–6.2 mm.

Labrum width smaller than clypeus width; proximal row of setae on dorsal face of labrum with < 30 setae; ventral side of labrum with stout setae in a row; maxillary palp segment III 1.7–2.2 × as long as base width; posterior margin of abdominal segment VIII with well-developed denticles.

Head. Yellowish, pale brown between ocelli and below compound eyes. Antenna yellowish. Mouth parts pale brown, mandibles medium brown. Thorax. Pronotum medium brown, dark brown laterally and posteriorly, as well as two elongated dots close to the median line. Mesonotum medium brown, dark brown laterally and on the anterolateral corners; costal and subcostal areas of forewing pads strongly tinted with black (Fig. 24B). Metanotum medium brown. Femur whitish, apex washed with pale orange. Tibia whitish, inner margin and apex washed with grey. Tarsi whitish. Abdomen. Terga brownish orange, with two longitudinal dark brown large stripes on segments I–IX; segment X medium brown. Sterna brownish orange, lateral margins washed with grey, posterolateral projections XIII–IX black. Gills purplish grey. Terminal filaments uniformly pale brown.

Head. Labrum cordiform (Fig. 25A). Ratio labrum width/clypeus width 0.86–0.97. Ratio labrum width/insertion width 1.26–1.30. Medial emargination shallow with flat denticles. Proximal row of setae present, simple. Number of setae on proximal row 19–23. Distal row of setae present, simple. Median part of labrum in ventral view with stout setae in row (Fig. 25B). Mandibles. Outer margin with middle and distal tuft of setae. Right mandible with 9–11 setae below the mola (Fig. 25C). Maxillae. Posterior margin of cardo with many hair-like setae and one stout and long seta in submarginal position (Fig. 34B). Proximo-lateral margin of stipes with a single stout and medium size seta. Apical-ventral row with 23–25 pectinate setae. Maxillary palp segment II 0.82–0.86 as long as segment I. Maxillary palp segment III 0.71–0.77 as long as segment I (Fig. 25D). Stout setae on outer margin of segment I absent. Segment II inner margin with two to five stout setae. Segment II outer margin with three or four stout setae. Segment III inner margin with 4–6 stout setae, dorsal surface with few setae at apex. Segment III 1.82–2.44 × as long as base width. Labium. Labial palp segment II 0.67–0.80 as long as segment I. Labial palp segment III 0.79–0.90 as long as segment I (Fig. 25E). Segment I inner margin with 5–9 stout setae, outer margin with 10–12 stout setae. Segment II outer margin with one or two stout setae. Segment III dorsal face with 5–7 stout setae. Segment III 3.50–3.80 × as long as base width. Hypopharynx. Apex of superlingua emarginated (Fig. 25F).

Thorax. Dorsal margin of femur with a row of long, stout, pointed setae; submarginal row with shorter setae; ventral margin with short, blunt, stout setae and a submarginal row of feathered setae; central area of upper surface without setae (Fig. 26A–B). Outer margin of fore tibia without setae; mid tibia with a row of stout, pointed setae on ventral margin; outer margin with only some hair-like setae; hind tibia covered with numerous simple and feathered setae, outer margin with a dense row of stout, pointed setae. Tarsal claw hooked, with 10–11 subequal denticles (Fig. 26C).

Abdomen. Gill I dorsal lamella narrow and lanceolate, ventral lamella shaped as other gills. Gills II–VII with dorsal and ventral lamellae of the same size. Gill II to VI with ~ 40 filaments on each lamella (Fig. 26G); gill VII with ~ 25 filaments on each lamella. Filamentous projections along entire margin. Posterior margin of terga VIII and IX with denticles (Fig. 26D–E). Posterior margin of terga VII with minute denticles (Fig. 26F). Posterior margin of terga V–VI without denticles. Posterolateral projections present on segment VII–IX.

This species is named after the Latin word iter, meaning path, to remember the new species has been collected along the trail from Port Moresby to Kokoda.

Holotype. • Papua New Guinea, nymph on slide, Central Province, Kokoda Trek, 980 m, I.2008, 09°15.933'S, 147°36.590'E, Posman col [PNG169], GBIFCH01223118 (ZSM). Paratype. • Papua New Guinea, 1 nymph in ethanol, GBIFCH01523481, same data as holotype (MZL).

Body length, female: 4.5 mm.

Labrum width equal to clypeus width; proximal row of setae on dorsal face of labrum with > 35 setae; maxillary palp segment III < 1.5 × as long as base width; tarsal claw with four teeth, distal one much larger than the others; posterior margin of abdominal segment VII–VIII without denticles; posterolateral projections of the abdomen present on segments VII–IX.

Head. Pale brown washed with grey, triangular yellowish area between front ocellus and clypeus. Mouth parts greyish brown. Thorax. Pronotum greyish brown, dark brown on lateral margins and along the sagittal line. Mesonotum pale brown washed with grey. Metanotum pale brown washed with grey. Femur upper surface greyish brown except apex yellowish, lower surface pale grey. Tibia yellowish brown, apex greyish. Tarsi yellowish to orange- brown. Abdomen. Terga more or less uniformly greyish brown, segments II–VII with a dark dot above the gill insertion, segments I–VIII with a paler sagittal line. Sterna yellowish grey in median part, segments I–VII greyish laterally, segment IX darker. Gills purplish grey. Terminal filaments uniformly pale brown.

Head. Labrum cordiform (Fig. 27A). Ratio labrum width/clypeus width 1.00. Ratio labrum width/insertion width 1.40. Medial emargination shallow with flat denticles. Proximal row of setae present simple. Number of setae on proximal row 42. Distal row of setae present, simple. Median part of labrum in ventral view with only scattered stout setae (Fig. 27B). Mandibles. Outer margin with row of setae on distal 1/2. Ending of the row near incisor. Right mandible with nine setae below the mola (Fig. 27C). Maxillae. Posterior margin of cardo broken. Proximo-lateral margin of stipes with two long and two short stout setae (Fig. 34F). Apical-ventral row with 24 pectinate setae. Maxillary palp segment II 0.81 as long as segment I. Maxillary palp segment III 0.80 as long as segment I (Fig. 27D). Stout setae on outer margin of segment I absent. Segment II inner margin with two stout setae. Segment II outer margin without stout setae. Segment III inner margin with three stout setae, dorsal surface with few setae at apex. Segment III 1.45 × as long as base width. Labium. Labial palp segment II 0.69 as long as segment I. Labial palp segment III 0.88 as long as segment I (Fig. 27E). Segment I inner margin with seven stout setae, outer margin with eight stout setae. Segment II outer margin with three stout setae. Segment III dorsal face with three stout setae. Segment III 3.0 × as long as base width. Hypopharynx. Apex of superlingua rounded (Fig. 27F).

Thorax. Dorsal margin of femur with a row of long, stout, pointed setae; submarginal row with setae of irregular length, as long or half the length of dorsal margin; ventral margin with short, blunt, stout setae and a submarginal irregular row of simple setae (Fig. 28A); central area of upper surface with few stout, pointed setae. Fore tibia covered with numerous simple and feathered setae on ventral margin; outer margin of fore tibia without setae; mid- and hind tibia with a row of stout, pointed setae on ventral margin, outer margin without setae on mid tibia, with a sparse row of stout and pointed setae on hind tibia. Tarsal claw hooked with 4 denticles progressively larger, except distal much larger (Fig. 28B).

Abdomen. Gill I dorsal lamella narrow and lanceolate, ventral lamella shaped as other gills. Gills II–VII with dorsal and ventral lamellae of the same size. Gill I with ~ 20 filaments on the ventral lamella. Gill II–VI with 25–30 filaments on each lamella. Posterior margin of terga IX with denticles. Posterior margin of terga V–VIII without denticles. Posterolateral projections present on segment VII–IX.

This species is named after our colleague and friend Dr Jean-Luc Gattolliat (Naturéum, Lausanne), in recognition of his outstanding contributions to the Baetidae in general and to those of New Guinea in peculiar.

Holotype • Indonesia, Papua, nymph on slide, Road Nabire–Enarotali KM 95, 160 m, 22.X.2011, 03°34.193'S, 135°49.246'E, M. Balke col [PAP13], GBIFCH01223119 (MZB). Paratypes. • Indonesia, Papua, one nymph on slide, GBIFCH01223150, 2 nymphs in ethanol, GBIFCH01523482, same data as holotype (MZL).

Body length, male: 5 mm.

Apical ventral row of maxilla composed of 25–35 pectinate setae; outer margin of maxillary palp II with five stout setae; outer margin of labial palp segment II with five stout setae; tarsal claw with six or seven teeth progressively larger; posterior margin of abdominal segment IX with well-developed denticles; posterolateral projections of the abdomen present on segments VI–IX.

Head. Pale brown, medium brown between ocelli. Antenna whitish. Compound eye lower portion black, upper portion reddish brown. Mouth parts pale brown. Thorax. Pronotum medium brown. Mesonotum medium brown, parapsidal sutures dark brown. Metanotum medium brown. Femur yellowish, apex with an orange to pale brown dot. Tibia: fore- and middle tibiae yellowish, hind tibia orange to brown. Tarsi whitish. Abdomen. Terga medium brown, with a pale and narrow sagittal line, segment IX lighter with two dark maculae, segment X greyish brown (Fig. 24C). Sterna pale brown, posterior margin of segments II to VIII medium brown. Gills with lamellae pale grey, filaments greyish purple. Terminal filaments yellowish, articulations of ~ 15 first segments tinted in brownish purple.

Head. Labrum cordiform (Fig. 29A). Ratio labrum width/clypeus width 0.93. Ratio labrum width/insertion width 1.29. Medial emargination shallow with flat denticles. Proximal row of setae present, simple. Number of setae on proximal row ~ 31. Distal row of setae present, simple. Median part of labrum in ventral view with stout setae in row (Fig. 29B). Mandibles. Outer margin with middle and distal tufts of setae. Ending of the row near incisor. Right mandible with ten setae below the mola (Fig. 29C). Maxillae. Posterior margin of cardo with numerous hair-like setae and few stout setae, with a long and stout seta in submarginal position. Proximo-lateral margin of stipes with a single stout and long seta (Fig. 34G). Apical-ventral row with 20 pectinate setae. Maxillary palp segment II 0.85 as long as segment I. Maxillary palp segment III 0.69 as long as segment I (Fig. 29D). Stout setae on outer margin of segment I absent. Segment II inner margin with five stout setae. Segment II outer margin with five stout setae. Segment III inner margin with four stout setae, dorsal surface with few setae at apex. Segment III 1.73 × as long as base width. Labium. Labial palp segment II 0.84 as long as segment I. Labial palp segment III 0.74 as long as segment I (Fig. 29E). Segment I inner margin with nine stout setae, outer margin with ten stout setae. Segment II outer margin with five stout setae. Segment III dorsal face with four stout setae. Segment III 3.11 × as long as base width. Hypopharynx. Apex of superlingua emarginated (Fig. 29F).

Thorax. Dorsal margin of femur with a row of long, stout, pointed setae; submarginal row with setae of irregular length, row as long or half the length of dorsal margin; ventral margin with short, blunt, stout setae and a submarginal irregular row of simple setae (Fig. 30A–B); central area of upper surface with few long, pointed setae. Fore tibia with three rows of numerous simple and feathered setae on ventral margin; outer margin of fore tibia with few hair-like setae; middle tibia with two rows of long, pointed setae on ventral margin, outer margin with numerous hair-like setae; hind tibia with a row of long, thin setae on ventral margin, with a sublateral row of simple and feathered setae, ventral surface covered with simple and feathered setae, outer margin with long, pointed stout setae. Tarsal claw hooked with six or seven denticles progressively larger apically (Fig. 30C).

Abdomen. Gill I dorsal lamella narrow and lanceolate, ventral lamella shaped as other gills. Gills II–VII with dorsal and ventral lamellae of the same size. Gill II–VI with ~ 25 filaments on each lamella, gill VII with ~ 15 filaments. Posterior margin of terga IX with denticles. Posterior margin of terga VIII with minute denticles. Posterior margin of terga V–VII without denticles. Posterolateral projections present on segments VI–IX.

This species is dedicated to Dr Pascale Derleth Sartori, wife of the first author.

Holotype. • Papua New Guinea, male nymph on slide, National Capital District, Varirata National Park, 600 m a.s.l., 09°26.13'S / 147°22.09'E, 16.XII.2007, Balke and Sagata col [PNG159], GBIFCH01223120 (ZSM). Paratypes. • Papua New Guinea, 4 nymphs in ethanol, GBIFCH01523483, 2 nymphs entirely mounted on slide, GBIFCH01223121, GBIFCH01223122, same data as holotype (MZL).

Body length, male: 4.5–4.8 mm; female: 4.9–5.2 mm.

Tarsal claw straight, without teeth; apical ventral row of maxilla composed of 16–20 pectinate setae; posterior margin of abdominal segment IX with minute denticles; posterolateral projections of the abdomen present on segments VIII–IX only.

Head. Yellowish brown, darker between ocelli. Triangular brown mark present on inner margin of compound eye. Antenna yellowish. Compound eye lower portion black, upper portion dark brown, not rounded but oblique and elongated. Mouth parts yellowish. Thorax. Pronotum yellowish washed with greyish brown laterally, irregular brown mark close to median line. Mesonotum yellowish washed with greyish laterally; distal half of costal field of fore wing pad orangish. Metanotum yellowish washed with greyish laterally. Femur yellowish white, dorsal surface apically washed with orange (Fig. 24A). Tibia whitish, washed with brown on dorsal surface. Tarsi yellowish white. Abdomen. Terga orangish with dark brown rectangular marks laterally on segments I to VI, subtriangular on segment VII and IX, segment VIII entirely dark brown. Sterna uniformly whitish becoming yellowish on the last segments. Gills purplish black. Terminal filaments yellowish white washed with orange proximally.

Head. Labrum cordiform (Fig. 31A). Ratio labrum width/clypeus width 0.89–0.98. Ratio labrum width/insertion width 1.22–1.24. Medial emargination shallow with flat denticles. Proximal row of setae present, simple. Number of setae on proximal row ~ 16–37. Distal row of setae present, simple. Median part of labrum in ventral view with stout setae in row (Fig. 31B). Mandibles. Outer margin with middle and distal tuft of setae. Right mandible with 6–11 setae below mola (Fig. 31C). Maxillae. Posterior margin of cardo with few hair-like setae and a long and stout seta in submarginal position. Proximo-lateral margin of stipes with a single stout and medium size seta. Apical-ventral row with 16–20 pectinate setae. Maxillary palp segment II 0.79–0.92 as long as segment I. Maxillary palp segment III 0.62–0.68 as long as segment I (Fig. 31D). Stout setae on outer margin of segment I absent. Segment II inner margin with two or three stout setae. Segment II outer margin without stout setae. Segment III inner margin with 2–4 stout setae, dorsal surface with few setae at apex. Segment III 1.5–2.0 × as long as base width. Labium. Labial palp segment II 0.72–0.76 as long as segment I. Labial palp segment III 0.79–0.92 as long as segment I. Segment I inner margin with 7–12 stout setae, outer margin with 8–12 stout setae. Segment II outer margin without or with one or two stout setae. Segment III dorsal face with five or six stout setae. Segment III 3.0–3.67 × as long as base width. Hypopharynx. Apex of superlingua emarginated (Fig. 31E).

Thorax. Dorsal margin of femur with a row of long, stout, pointed setae; incomplete submarginal row, sometimes with scattered long, pointed setae; ventral margin with short, blunt, stout setae; central area of upper surface without setae (Fig. 32A–B). Inner margin of fore- and mid- tibia covered with several rows of simple and feathered setae; inner margin of hind tibia with several rows of simple and feathered setae; outer margin of fore- and mid- tibia naked, with a row of stout, pointed setae on hind tibia, especially developed in the distal half. Tarsal claw straight. Denticles absent (Fig. 32C).

Abdomen. Gill I dorsal lamella narrow and lanceolate, ventral lamella shaped as other gills (Fig. 32E). Gills II–VII with dorsal and ventral lamellae of the same size. Ventral lamella of gill I with ~ 25 long filaments; dorsal and ventral lamellae of gills II–VI with ~ 40 very long filaments each (Fig. 32F); gill VII with ~ 20 filaments on each lamella. Posterior margin of terga IX with small denticles (Fig. 32D). Posterior margin of terga V–VIII without denticles. Posterolateral projections on segments VIII–IX.

Eggs. General shape elongated. Size ca 185 μm × 80 μm. Chorionic surface with irregular longitudinal ridges connected transversally, forming rectangular smooth areas in-between (Fig. 64A). On each pole arise basally six equally distant pairs of long, apically hooked attachment structure (Fig. 64B). Micropyle not visible.

This species is dedicated to Samuel Sartori, grandson of the first author.

• Indonesia, Papua, one nymph on slide, Road Nabire–Enarotali KM 52, 555 m, 23.X.2011, 03°30.107'S, 135°42.971'E, M. Balke col [PAP17], GBIFCH01223123 (MZL).

Nymph. Labrum width smaller than clypeus width; proximal row of setae on dorsal face of labrum with < 30 setae; ventral side of labrum with scattered stout setae; maxillary palp segment III ~ 1.7 × as long as base width; posterior margin of abdominal segment VIII with minute denticles; posterolateral projections on the abdomen present on segments VII–IX.

Head. Labrum cordiform (Fig. 33A). Ratio labrum width/clypeus width 0.95. Ratio labrum width/insertion width 1.33. Medial emargination shallow with flat denticles. Proximal row of setae present simple. Number of setae on proximal row 23. Distal row of setae present, simple. Median part of labrum in ventral view with only scattered stout setae (Fig. 33B). Mandibles. Outer margin with middle and distal tuft of setae. Right mandible with 11 setae below the mola (Fig. 33C). Maxillae. Posterior margin of cardo with few hair-like setae. Proximo-lateral margin of stipes with two long and two short stout setae. Apical-ventral row with 24 pectinate setae. Maxillary palp segment II 0.78 as long as segment I. Maxillary palp segment III 0.74 as long as segment I (Fig. 33D). Stout setae on outer margin of segment I absent. Segment II inner margin with three stout setae. Segment II outer margin with four stout setae. Segment III inner margin with three stout setae, dorsal surface with few setae at apex. Segment III 1.70 × as long as base width. Labium. Labial palp segment II 0.71 as long as segment I. Labial palp segment III 0.90 as long as segment I (Fig. 33E). Segment I inner margin with seven stout setae, outer margin with ten stout setae. Segment II outer margin with one stout seta. Segment III dorsal face with five stout setae. Segment III 3.00 × as long as base width. Hypopharynx. Apex of superlingua emarginated (Fig. 33F).

Abdomen. Gill I dorsal lamella narrow and lanceolate, ventral lamella shaped as other gills. Gill I with ~ 18 filaments. Posterior margin of terga IX with denticles. Posterior margin of terga VII–VIII with minute denticles. Posterior margin of terga V–VI without denticles. Posterolateral projections present on segments VII–IX.

Ventral lamella of gills highly reduced or absent, ventral lamella of first gill with < 8 processes, even absent in some species; claw with generally 3–6 teeth, rarely up to 8; posterior margin of abdominal segment VIII generally without denticles; posterolateral expansions on abdominal segments VIII–IX, rarely VII–IX.

Nonnullidens alvarezi sp. nov. [Papua New Guinea], N. anga sp. nov. [Papua New Guinea], N. billhilli Grant & Peters, 1993 [Papua New Guinea, New Britain], N. boonsoongi sp. nov. [Papua New Guinea], N. cozzarolae sp. nov. [Papua New Guinea], N. depapricus Kluge, 2013 [Indonesia, Papua], N. fuyugensis sp. nov. [Papua New Guinea], N. hsui (Peters & Tsui, 1972) [Papua New Guinea], N. kaltenbachi sp. nov. [Papua New Guinea], N. marcelae sp. nov. [Indonesia, Papua], N. mariae (Peters & Tsui, 1972) [Papua New Guinea], N. moiorum sp. nov. [Indonesia, Papua], N. niger Kluge, 2013 [Indonesia, Papua], N. reductus Kluge, 2013 [Indonesia, Papua], N. silvaepumilorum sp. nov. [Papua New Guinea], N. variegatus Kluge, 2013 [Indonesia, Papua]

Species of the genus Nonnullidens are restricted to New Guinea; N. billhilli is also reported from New Britain (Grant and Peters 1993).

Holotype. • Papua New Guinea, one nymph on slide, Eastern Highlands Province, Akameku – Brahmin, Bismarck Range, 2200 m, 23.XI.2006, 05°56.801'S, 145°22.238'E, M. Balke & Kinibel col [PNG106], GBIFCH01223124 (ZSM). Paratypes. • Papua New Guinea, 2 nymphs in ethanol (UFVB), 2 nymphs on slide, GBIFCH01223125, GBIFCH01223126, 7 nymphs in ethanol, GBIFCH01523456 same data as holotype (MZL). • Papua New Guinea, one nymph on slide, GBIFCH01223127, gills and legs in ethanol, GBIFCH01523457, Gulf Province, Marawaka, Mala, 1400 m, 11.XI.2006, 07°05.664'S, 145°44.467'E, M. Balke & Kinibel col [PNG90] (MZL).

Body length, male: 5.5 mm; female: 6.5 mm.

Femora uniformly dark brown, without transverse bands; tarsal claw with 6–8 teeth; gill I–II uni-lamellate, without ventral lamella; gills III–VI with 5–8 filaments on dorsal lamella and four or five filaments on ventral lamella, gill VII with four or five filaments on dorsal lamella and one or two filaments on ventral lamella.

Head. Vertex dark brown, frons medium brown with an oval pale brown macula near the frons ocellus. Antenna broken. Compound eye upper portion reddish brown. Mouth parts pale brown. Thorax. Pronotum medium brown, lateral margins dark brown as well as medio anteriorly. Mesonotum pale brown washed with dark brown as in Fig. 36B. Femur greyish brown, except the base whitish. Tibia yellowish grey, apex of fore tibia blackish. Tarsi pale brown. Abdomen. Terga dark brown, with a thin sagittal line lighter on segments V–VII, becoming wider on segments VIII–X, less visible on mature nymphs. Sterna greyish brown, segments VIII–IX darker. Gills greyish purple. Terminal filaments yellowish.

Head. Labrum cordiform (Fig. 37A). Ratio labrum width/clypeus width 0.99–1.10. Ratio labrum width/insertion width 1.35–1.53. Medial emargination shallow with flat denticles. Proximal row of setae present, simple. Number of setae on proximal row ~ 45–61. Distal row of setae present, simple. Median part of labrum in ventral view with scattered thin setae (Fig. 37B). Mandibles. Outer margin with row of setae on distal 1/2. Ending of the row near incisor. Right mandible with 6–10 setae below the mola (Fig. 37C). Maxillae. Posterior margin of cardo with few hair-like and stout setae, together with one long and stout seta in submarginal position. Proximo-lateral margin of stipes with a bunch of six stout setae increasing in size distally. Apical-ventral row with 21–24 pectinate setae. Maxillary palp segment II 0.83–0.91 as long as segment I. Maxillary palp segment III 0.67–0.82 as long as segment I (Fig. 37D). Stout setae on outer margin of segment I absent. Segment II inner margin with 2–4 stout setae. Segment II outer margin with 1–4 stout setae. Segment III inner margin with 4–6 stout setae, dorsal surface with few setae at apex. Segment III 1.83–2.00 × as long as base width. Labium. Labial palp segment II 0.72–0.80 as long as segment I. Labial palp segment III 0.78–0.88 as long as segment I (Fig. 37E). Segment I inner margin with 10–15 stout setae, outer margin with 9–12 stout setae. Segment II outer margin with four or five stout setae. Segment III dorsal face with 3–5 stout setae. Segment III 3.00–3.86 × as long as base width. Hypopharynx. Apex of superlingua truncate (Fig. 37F).

Thorax. Dorsal margin of femur with a row of very long, stout, pointed setae; an irregular submarginal row of pointed setae as long as or a little bit shorter than those on the dorsal margin on fore femur, more regular on mid- and hind femur; ventral margin with short, stout setae and an irregular submarginal row of long, pointed stout setae; central area of upper surface with few long and short, stout, pointed setae (Fig. 38A–B). Fore tibia with two rows of long simple or feathered setae on ventral margin, dorsal margin with few hair-like setae; middle tibia with a row of short, simple, pointed setae on ventral margin and a submarginal row of long, simple setae, outer margin with numerous hair-like setae; hind tibia with two rows of simple and feathered long, stout setae on ventral margin, outer margin with short and long pointed stout setae and numerous hair-like setae. Tarsal claw slightly hooked with 6–8 denticles progressively larger distally (Fig. 38C).

Abdomen. Gills I and II with single lamella (Fig. 38E–F). Gills III–VII with dorsal and ventral lamellae, ventral lamella smaller. Gill I with three filaments on dorsal lamella, without ventral lamella, gill II with four or five filaments on dorsal lamella, without ventral lamella, gills III–VI with 5–8 filaments on dorsal lamella and four or five on ventral lamella (Fig. 38G), gill VII with four or five filaments on dorsal lamella and one or two filaments on ventral lamella (Fig. 38H). Posterior margin of terga IX with minute denticles (Fig. 38D). Posterior margin of terga V–VIII without denticles. Posterolateral projections present on segments VIII–IX.

This species is dedicated to Prof. Nadir Alvarez, current director of Naturéum and instigator of the Cozzarolo et al. paper who revealed the diversity of New Guinean Thraulini.

Holotype. • Papua New Guinea, one nymph on slide, Morobe Province, Menyamya, Mt Inji, 1700 m, 14.XI.2006, approx. 07°14.813'S, 146°01.330'E, M. Balke & Kinibel col [PNG96], GBIFCH01223129 (ZSM). Paratype. • Papua New Guinea, one nymph in ethanol, GBIFCH01523458, same data as holotype (MZL).

Body length, female: 4.8 mm.

Labrum cordiform; dorsal face of labrum with proximal and distal rows multiple; femora pale brown with two transverse dark brown bands in middle and at apex; gill I with ventral lamella composed of a single filament; gill II–VI with six or seven filaments on dorsal lamella and two or three on ventral lamella; gill VII with four or five filaments on dorsal lamella and a single one on ventral lamella.

Head. Vertex greyish brown, with characteristic yellowish maculae posteriorly, frons greyish brown with a middle pale brown oval macula. Antenna: scape greyish, pedicel and flagellum broken. Mouth parts pale brown. Thorax. Pronotum pale brown, washed with medium brown laterally and medio-anteriorly. Mesonotum pale brown, washed with medium brown as in Fig. 35A. Femur yellowish brown, with a large median greyish brown stripe and another one at apex. Tibia yellowish. Tarsi pale brown. Abdomen. Terga dark brown, segments VII–X somewhat lighter, pale brown along the sagittal line. Sterna yellowish grey, yellowish on lateral margins. Gills greyish brown. Terminal filaments yellowish, basally yellowish brown.

Head. Labrum cordiform (Fig. 39A). Ratio labrum width/clypeus width 1.05. Ratio labrum width/insertion width 1.36. Medial emargination narrow with flat denticles. Proximal row of setae present, multiple. Number of setae on proximal row ~ 104. Distal row of setae multiple. Median part of labrum in ventral view with scattered thin setae (Fig. 39B). Mandibles. Outer margin with row of setae on distal 1/2. Ending of the row near incisor. Right mandible with seven setae below the mola (Fig. 39C). Maxillae. Posterior margin of cardo with many long hair-like setae. Proximo-lateral margin of stipes twisted. Apical-ventral row with 23 pectinate setae. Maxillary palp segment II 0.88 as long as segment I. Maxillary palp segment III 0.68 as long as segment I (Fig. 39D). Stout setae on outer margin of segment I absent. Segment III dorsal surface half covered with setae. Segment III 1.55 × as long as base width. Labium. Labial palp segment II 0.82 as long as segment I. Labial palp segment III 0.71 as long as segment I (Fig. 39E). Segment I inner margin with nine stout setae, outer margin with 14 stout setae. Segment II outer margin with four stout setae. Segment III dorsal face with four stout setae. Segment III 2.67 × as long as base width. Hypopharynx. Apex of superlingua truncate.

Thorax. Dorsal margin of femur with a row of very long, thin, pointed setae; regular submarginal row of pointed setae as long as those on the dorsal margin; ventral margin with relatively long, pointed setae and an irregular submarginal row of long, pointed thin setae (Fig. 40A–B); central area of upper surface with very long pointed setae, as long as those on the dorsal margin. Fore tibia with two or three rows of long simple or feathered setae on ventral margin, dorsal margin with few hair-like setae; middle tibia with a row of short, simple, pointed setae on ventral margin and a submarginal row of long, simple setae, outer margin with numerous hair-like setae; hind tibia with a marginal row of simple short, thin setae and a submarginal row of long to very long setae on ventral margin, outer margin with long to very long pointed setae and hair-like setae (Fig. 40C). Tarsal claw hooked with three or four denticles progressively larger, except distal much larger.

Abdomen. Gill I with both lamellae. Gills II–VII with dorsal and ventral lamellae, ventral lamella smaller. Gill I with three filaments on dorsal lamella, ventral lamella with a single filament (Fig. 40E); gill II–VI with six or seven filaments on dorsal lamella and two or three on ventral lamella (Fig. 40F); gill VII with four or five filaments on dorsal lamella and a single one on ventral lamella (Fig. 40G). Posterior margin of terga IX with minute denticles (Fig. 40D). Posterior margin of terga V–VIII without denticles. Posterolateral projections present on segments VIII–IX.

The specific name is to recognize the Anga people who live in the area where the new species has been found and is a noun in apposition.

Holotype. • Papua New Guinea, one nymph on slide, Central Province, Kokoda Trek, 1400 m, I. 2008, 09°01.952'S, 147°44.455'E, Posman col [PNG172], GBIFCG01223130 (ZSM). Paratypes. two nymphs on slide, GBIFCH01223131, GBIFCH01223132, 10 nymphs in ethanol, GBIFCH01523459, same data as holotype (MZL). Other material. • Papua New Guinea, one nymph on slide, GBIFCH01223133, gills and legs in ethanol, GBIFCH01523460, Western Highlands, Simbai area, 2500 m, 8. III.2007, 05°14.202'S, 144°33.651'E, Kinibel col [PNG150] (MZL).

Body length, male: 5 mm; female: 5.5 mm.

Femora uniformly dark brown, without transverse bands; tarsal claw with 6–8 teeth; gill I and II uni-lamellate, without ventral lamella; gills III–VI with five or six filaments on dorsal lamella and a single or two filaments on ventral lamella, gill VII with three filaments on dorsal lamella and a single filament on ventral lamella.

Head. Vertex dark brown, with characteristic yellowish maculae posteriorly, pale brown between lateral ocelli and eyes, frons medium brown with a middle pale brown oval macula. Antenna broken. Compound eye upper portion reddish brown. Mouth parts pale brown. Thorax. Pronotum medium brown, dark brown laterally and anteromedially as in Fig. 36A. Mesonotum pale brown washed with dark brown. Femur greyish brown, whitish at base. Tibia yellowish, apex greyish brown on fore- and mid-tibia. Tarsi yellowish brown. Abdomen. Terga dark brown, paler along the sagittal line on segments VII–IX. Sterna pale brown, greyish brown in the middle, pale brown on each side of a sagittal line. Gills greyish brown. Terminal filaments yellowish.

Head. Labrum cordiform (Fig. 41A). Ratio labrum width/clypeus width 0.99–1.04. Ratio labrum width/insertion width 1.36–1.45. Medial emargination shallow with flat denticles. Proximal row of setae present, simple. Number of setae on proximal row ~ 42–69. Distal row of setae present, simple. Median part of labrum in ventral view with scattered thin setae (Fig. 41B). Mandibles. Outer margin with row of setae on distal 1/2. Ending of the row near incisor. Right mandible with eight or nine setae below the mola (Fig. 41C). Maxillae. Posterior margin of cardo with few hair-like setae, together with two stout and long setae in submarginal position. Proximo-lateral margin of stipes with a bunch of five stout setae of irregular length. Apical-ventral row with 22–24 pectinate setae. Maxillary palp segment II 0.86–1.00 as long as segment I. Maxillary palp segment III 0.64–0.95 as long as segment I (Fig. 41D). Stout setae on outer margin of segment I absent. Segment II inner margin with two or three stout setae. Segment II outer margin without stout setae. Segment III inner margin with four stout setae, dorsal surface with few setae at apex. Segment III 1.67–2.00 × as long as base width. Labium. Labial palp segment II 0.70–0.80 as long as segment I. Labial palp segment III 0.74–0.80 as long as segment I (Fig. 41E). Segment I inner margin with 13–15 stout setae, outer margin with 12–15 stout setae. Segment II outer margin with three or four stout setae. Segment III dorsal face with 3–5 stout setae. Segment III 2.91–3.30 × as long as base width. Hypopharynx. Apex of superlingua truncate (Fig. 41F).

Thorax. Dorsal margin of femur with few long, stout, pointed setae; an irregular submarginal row of pointed setae as long as or a little bit shorter than those on the dorsal margin; ventral margin with short, stout setae and an irregular submarginal row of long, pointed stout setae; central area of upper surface with numerous long and short, stout, pointed setae (Fig. 42A). Fore tibia with several rows of long simple or feathered setae on ventral margin, dorsal margin with few hair-like setae; middle tibia with a row of short, simple, pointed setae on ventral margin and a submarginal row of long, simple setae, dorsal margin with few hair-like setae; hind tibia with two rows of simple and feathered long, stout setae on ventral margin, dorsal margin with short and long pointed stout setae and few hair-like setae. Tarsal claw slightly hooked with seven or eight denticles progressively larger apically (Fig. 42B).

Abdomen. Gill I–II with single lamella. Gills III–VII with dorsal and ventral lamellae, ventral lamella smaller. Gill I uni-lamellate, with two or three filaments (Fig. 42D), gill II uni-lamellate with four filaments (Fig. 42E), gills III–VI with five or six filaments on dorsal lamella and a single or two filaments on ventral lamella (Fig. 42F), gill VII with three filaments on dorsal lamella and a single filament on ventral lamella (Fig. 42G). Posterior margin of terga IX with minute denticles (Fig. 42C). Posterior margin of terga V–VIII without denticles. Posterolateral projections present on segments VIII–IX.

This species is dedicated to Prof. Boonsatien Boonsoong (Kasetsart University, Bangkok, Thailand) for his outstanding contributions to our knowledge of Southeast Asian Ephemeroptera.

Holotype. • Papua New Guinea, one nymph on slide, Eastern Highlands Province, Marawaka Ande, 1700–1800 m, 9.XI.2006, approx. 07°01.697'S, 145°49.807'E, M. Balke & Kinibel col [PNG87], GBIFCH01223134 (ZSM). Paratypes. • Papua New Guinea, 2 nymphs on slide, GBIFCH01223135, GBIFCH01223136, 5 nymphs in ethanol, GBIFCH01523461, same data as holotype (MZL). • Papua New Guinea, 3 nymphs on slide, GBIFCH01223137, GBIFCH01223138, GBIFCH01223139, one nymph in ethanol, GBIFCH01523462, Central Province, Kokoda Trek, 1390 m, I. 2008, 09°00.338'S, 147°44.252'E, Posman col [PNG173] (MZL). • Papua New Guinea, one nymph on slide GBIFCH01223140, 2 nymphs in ethanol, GBIFCH01523463 Western Highlands Province, Kundum, 1400 m, 3.III.2007, 05°16.096'S, 144°27.869'E, Kinibel col [PNG142] (MZL).

Body length, male: 5 mm; female: 5.25 mm.

Femora uniformly dark brown, without transverse bands; gill I with ventral lamella composed of one or two filaments; gills II–VI with four to six filaments on dorsal lamella and three or four on ventral lamella, gill VII with 2–4 filaments on each lamella; posterolateral projections present on segments VII–IX.

Head. Vertex greyish brown, with characteristic yellowish maculae posteriorly, frons medium brown with a middle pale brown oval macula. Antenna: scape yellowish, pedicel and flagellum pale brown. Compound eye upper portion reddish brown. Mouth parts pale brown. Thorax. Pronotum pale brown, dark brown laterally and medio-anteriorly. Mesonotum greyish brown laterally, with a pale brown central area as in Fig. 35C. Femur greyish brown, lighter at base. Fore tibia greyish brown, mid- and hind tibia pale brown, apex greyish brown. Tarsi pale brown. Abdomen. Terga dark brown, with a pale brown macula of the middle of segments IV–IX, increasing in size posteriorly, segment X almost completely pale brown. Sterna pale brown, greyish brown in the middle on each side of a sagittal line pale brown. Gills greyish brown. Terminal filaments yellowish to pale brown.

Head. Labrum cordiform (Fig. 43A). Ratio labrum width/clypeus width 1.06–1.13. Ratio labrum width/insertion width 1.46–1.56. Medial emargination shallow with flat denticles. Proximal row of setae present, simple. Number of setae on proximal row ~ 33–58. Distal row of setae present, simple. Median part of labrum in ventral view with only scattered stout setae (Fig. 43B). Mandibles. Outer margin with row of setae on distal 1/2. Ending of the row near incisor. Right mandible with 7–10 setae below the mola (Fig. 43C). Maxillae. Posterior margin of cardo with few hair-like and some stout setae, together with one long and stout seta in submarginal position. Proximo-lateral margin of stipes with a bunch of ten stout medium size and short setae. Apical-ventral row with 22–29 pectinate setae. Maxillary palp segment II 0.80–1.00 as long as segment I. Maxillary palp segment III 0.68–0.86 as long as segment I (Fig. 43D). Stout setae on outer margin of segment I absent. Segment II inner margin with four or five stout setae. Segment II outer margin without or with one stout seta. Segment III inner margin with 3–5 stout setae, dorsal surface with few setae at apex. Segment III 1.57–1.83 × as long as base width. Labium. Labial palp segment II 0.71–0.94 as long as segment I. Labial palp segment III 0.73–0.81 as long as segment I (Fig. 43E). Segment I inner margin with 10–12 stout setae, outer margin with 10–12 stout setae. Segment II outer margin with 3–5 stout setae, dorsal face with 3–5 stout setae. Segment III 2.90–3.40 × as long as base width. Hypopharynx. Apex of superlingua truncate (Fig. 43F).

Thorax. Dorsal margin of femur with a row of long, stout, pointed setae; a regular submarginal row of pointed setae as long as or a little bit shorter than those on the dorsal margin; ventral margin with short, stout setae and an irregular submarginal row of long, pointed stout setae; central area of upper surface with numerous long and short, stout, pointed setae (Fig. 44A–B). Fore tibia with several rows of long simple or feathered setae on ventral margin, dorsal margin with numerous hair-like setae; middle tibia with a row of short, simple, pointed setae on ventral margin and a submarginal row of long, simple setae, dorsal margin with numerous hair-like setae; hind tibia with two rows of simple long, stout setae on ventral margin, dorsal margin with short and long pointed stout setae and numerous hair-like setae. Tarsal claw slightly hooked with 4–6 denticles progressively larger apically (Fig. 44C).

Abdomen. Gill I with both lamellae. Gills II–VII with dorsal and ventral lamellae, ventral lamella smaller. Gill I with three or four filaments on dorsal lamella, ventral lamella with a single or two filaments (Fig. 44E), gills II–VI with 4–6 filaments on dorsal lamella and three or four on ventral lamella (Fig. 44F), gill VII with 2–4 filaments on each lamella (Fig. 44G). Posterior margin of terga IX with minute denticles (Fig. 44D). Posterior margin of terga V–VIII without denticles. Posterolateral projections present on segments VII–IX.

This species is dedicated to Dr Camille-Sophie Cozzarolo, who performed the first genetic work which revealed the diversity of New Guinean Thraulini.

Holotype. • Papua New Guinea, one nymph on slide, Central Province, Woitape, 1700 m, I. 2008, 08°31.290'S, 147°13.684'E, Posman col [PNG166], GBIFCH01223141 (ZSM). Paratypes. • Papua New Guinea, one nymph on slide GBIFCH01223142, 4 nymphs in ethanol, GBIFCH01523464, same data as holotype (MZL).

Body length, female: 4.5–5.0 mm.

Femora uniformly dark brown, without transverse bands; gill I uni-lamellate, without ventral lamella; gills II–VI with four or five filaments on dorsal lamella and two filaments on ventral lamella, gill VII with three or four filaments on dorsal lamella and a single one on ventral lamella.

Head. Vertex greyish brown, with characteristic yellowish maculae posteriorly, pale brown between lateral ocelli and eyes, frons medium brown with a middle pale brown oval macula. Antenna broken. Mouth parts pale brown. Thorax. Pronotum medium brown, greyish brown laterally and antero-medially. Mesonotum pale brown, washed with greyish brown laterally (Fig. 35B). Femur greyish brown, whitish at base. Tibia yellowish to pale brown. Tarsi greyish brown. Abdomen. Terga dark brown, with a paler sagittal line increasing in width on segments VIII–X. Sterna yellowish brown to greyish brown. Gills greyish purple. Terminal filaments yellowish.

Head. Labrum cordiform (Fig. 45A). Ratio labrum width/clypeus width 1.04–1.10. Ratio labrum width/insertion width 1.45–1.53. Medial emargination shallow with flat denticles. Proximal row of setae present, simple. Number of setae on proximal row ~ 36–37. Distal row of setae present, simple. Median part of labrum in ventral view with scattered thin setae (Fig. 45B). Mandibles. Outer margin with row of setae on distal 1/2. Ending of the row near incisor. Right mandible with 7–9 setae below the mola (Fig. 45C). Maxillae. Posterior margin of cardo with few hair-like and some stout setae (Fig. 34C). Proximo-lateral margin of stipes with a bunch of six stout medium size and long setae. Apical-ventral row with 23–27 pectinate setae. Maxillary palp segment II 0.79–0.80 as long as segment I. Maxillary palp segment III 0.79–0.80 as long as segment I (Fig. 45D). Stout setae on outer margin of segment I absent. Segment II inner margin with three or four stout setae. Segment II outer margin without stout setae. Segment III inner margin with five stout setae, dorsal surface with few setae at apex. Segment III 1.69–2.10 × as long as base width. Labium. Labial palp segment II 0.84–0.86 as long as segment I. Labial palp segment III 0.81–0.84 as long as segment I (Fig. 45E). Segment I inner margin with 11–14 stout setae, outer margin with 14 stout setae. Segment II outer margin with four or five stout setae. Segment III dorsal face with three or four stout setae. Segment III 3.20–3.40 × as long as base width. Hypopharynx. Apex of superlingua truncate (Fig. 45F).

Thorax. Dorsal margin of femur with a row of long, stout, pointed setae; a regular submarginal row of pointed setae as long as or a little bit shorter than those on the dorsal margin, shorter on hind femur; ventral margin with short, stout setae and an irregular submarginal row of long, pointed stout setae; central area of upper surface with numerous long and short, stout, pointed setae (Fig. 46A). Fore tibia with several rows of long simple or feathered setae on ventral margin, dorsal margin with numerous hair-like setae; middle tibia with a row of short, simple, pointed setae on ventral margin and a submarginal row of long, simple setae, dorsal margin with numerous hair-like setae; hind tibia with two rows of simple and feathered long, stout setae on ventral margin, dorsal margin with short and long pointed stout setae and numerous hair-like setae. Tarsal claw slightly hooked with four or five denticles progressively larger apically (Fig. 46B).

Abdomen. Gill I with single lamella. Gills II–VII with dorsal and ventral lamellae, ventral lamella smaller. Gill I uni-lamellate with three or four filaments (Fig. 46D), gills II–VI with four or five filaments on dorsal lamella and two filaments on ventral lamella (Fig. 46E), gill VII with three or four filaments on dorsal lamella and a single one on ventral lamella (Fig. 46F). Posterior margin of terga IX with minute denticles (Fig. 46C). Posterior margin of terga V–VIII without denticles. Posterolateral projections present on segments VIII–IX.

Eggs. General shape rounded. Size 130 μm × 80 μm. Chorionic surface smooth without visible attachment structures (Fig. 64C). Micropyle in equatorial position (Fig. 64D).

The specific name honours the tribes of the Fuyug (Fuyughe) people, who live where the new species has been found.

Holotype. • Papua New Guinea, one nymph on slide, Madang Province, Highway near Madang, ford, 80 m, 2–3.XII.2006, 05°24.405'S, 145°38.213'E, Binatang Boys col [PNG117], GBIFCH01223143, some gills and legs in ethanol, GBIFCH01523465 (ZSM). Paratype. • Papua New Guinea, one nymph on slide, GBIFCH01223144, Madang Province, Keki, Adalbert Mts, 400 m, 22.XI.2006, 04°43.058'S, 145°24.437'E, Binatang Boys col [PNG119] (MZL).

Nymph. Labrum emargination narrow and very deep with flat denticles; femora pale brown with two transverse dark brown bands in middle and at apex; central area of upper surface of femora with very few long and pointed setae; posterior margin of abdominal segments IX with minute denticles; posterolateral expansions on the abdomen present on segments VIII–IX.

Forefemur yellowish, washed with greyish brown in distal half, mid- and hind femur yellowish with a brownish band and in the middle and at apex. Tibia yellowish. Tarsi yellowish.

Head. Labrum cordiform (Fig. 47A). Ratio labrum width/clypeus width 1.03–1.07. Ratio labrum width/insertion width 1.40–1.42. Medial emargination deep, narrow with flat denticles. Proximal row of setae present, simple, shifted to distal margin. Number of setae on proximal row ~ 69–75. Distal row of setae present, simple. Median part of labrum in ventral view with scattered thin setae (Fig. 47B). Mandibles. Outer margin with row of setae on distal 1/2. Ending of the row near incisor. Right mandible with 4–6 setae below the mola (Fig. 47C). Maxillae. Posterior margin of cardo with few hair-like setae, together with one stout and long seta in submarginal position. Proximo-lateral margin of stipes with a bunch of six stout setae increasing in size distally, most distal much longer than the others. Apical-ventral row with 23–26 pectinate setae. Maxillary palp segment II 0.79–0.84 as long as segment I. Maxillary palp segment III 0.61 as long as segment I (Fig. 47D). Stout setae on outer margin of segment I absent. Segment II inner margin with one stout seta. Segment II outer margin without or with up to two stout setae. Segment III inner margin without or with up to three stout setae, dorsal surface half covered with setae. Segment III 1.25–1.50 × as long as base width. Labium. Labial palp segment II 0.68–0.86 as long as segment I. Labial palp segment III 0.58–0.66 as long as segment I (Fig. 47E). Segment I inner margin with 6–10 stout setae outer margin with 9–13 stout setae. Segment II outer margin with two or three stout setae, dorsal face with two or three stout setae. Segment III 2.71–2.75 × as long as base width. Hypopharynx. Apex of superlingua truncate (Fig. 47F).

Thorax. Dorsal margin of femur with a row of long, pointed setae; irregular submarginal row of long, pointed setae as long as those on the dorsal margin; ventral margin with short, pointed setae and an irregular submarginal row of longer and blunt setae (Fig. 48A–B); central area of upper surface with very few long, pointed setae (Fig. 48C). Fore tibia with several rows of long simple or feathered setae on ventral margin, dorsal margin with few hair-like setae; middle tibia with one row of short, simple, pointed setae, and an irregular submarginal row of long, pointed setae on ventral margin, dorsal margin with numerous hair-like setae; hind tibia with a row of short simple setae on ventral margin and a submarginal row of long, simple, pointed setae, outer margin with long and short pointed setae and numerous hair-like setae. Tarsal claw hooked with three denticles progressively larger apically (Fig. 48D).

Abdomen. Gill I with both lamellae. Gills II–VII with dorsal and ventral lamellae, ventral lamella smaller. Gill I with five filaments on dorsal lamella and a single filament on ventral lamella (Fig. 48E), gills II–VI with seven or eight filaments on dorsal lamella and three or four filaments on ventral lamella (Fig. 48F), gill VII with seven filaments on dorsal lamella and two filaments on ventral lamella (Fig. 48G). Posterior margin of terga IX with minute denticles. Posterior margin of terga V–VIII without denticles. Posterolateral projections present on segments VIII–IX.

This species is dedicated to Dr Thomas Kaltenbach (Naturéum, Lausanne) who revealed the megadiversity of the mayfly genus Labiobaetis in New Guinea.

Holotype. • Indonesia, Papua, one nymph on slide, Sorong Island, 95 m, 19.II.2006, 00°49.354'S, 131°24.196'E, M. Balke & Tindige col [BH20], GBIFCH01223151 (MZB). Paratypes. • Indonesia, Papua, one nymph on slide, GBIFCH01223152, one nymph in ethanol, GBIFCH01523466, same data as holotype (MZL).

Body length, female: 3.5 mm.

Labrum emargination narrow without denticles; central area of upper surface of femora with very few short blunt setae; femora pale brown with two transverse dark brown bands in middle and at apex; posterior margin of abdominal segments IX with well-developed denticles; posterolateral expansions on the abdomen present on segments VIII–IX.

Head. Vertex greyish brown, clypeus pale brown. Antenna broken. Mouth parts pale brown. Thorax. Pronotum pale brown, greyish brown on lateral margins, a greyish brown triangular macula in antero median position. Mesonotum pale brown, washed with greyish brown posteriorly (Fig. 50C). Femur yellowish, washed with pale brown in median area and at apex. Tibia uniformly yellowish. Tarsi yellowish. Abdomen. Terga uniformly greyish brown, posterior margin of segments IX and X pale brown. Sterna yellowish, sterna I–VII washed with greyish brown laterally. Gills greyish brown, filaments pale grey. Terminal filaments yellowish.

Head. Labrum rounded (Fig. 49A). Ratio labrum width/clypeus width 0.97–1.00. Ratio labrum width/insertion width 1.19–1.24. Medial emargination narrow without denticles. Proximal row of setae present, simple. Number of setae on proximal row ~ 51–75. Distal row of setae present, simple. Median part of labrum in ventral view with scattered thin setae (Fig. 49B). Mandibles. Outer margin with row of setae on distal 1/2. Ending of the row before incisor. Right mandible with 8–12 setae below the mola (Fig. 49C). Maxillae. Posterior margin of cardo with few hair-like setae, together with one stout and long seta in submarginal position. Proximo-lateral margin of stipes with a single stout and long seta. Apical-ventral row with 23–25 pectinate setae. Maxillary palp segment II 0.76–0.91 as long as segment I. Maxillary palp segment III 0.62–0.73 as long as segment I (Fig. 49D). Stout setae on outer margin of segment I absent. Segment II inner margin with three stout setae, outer margin without stout setae. Segment III inner margin with three or four stout setae, dorsal surface with few setae at apex, or half covered with setae. Segment III 1.78–2.05 × as long as base width. Labium. Labial palp segment II 0.75 as long as segment I. Labial palp segment III 0.56 as long as segment I (Fig. 49E). Segment I inner margin with ten stout setae, outer margin with 13 stout setae. Segment II outer margin without stout setae dorsal face with three stout setae. Segment III 2.71 × as long as base width. Hypopharynx. Apex of superlingua somewhat pointed (Fig. 49F).

Thorax. Dorsal margin of femur with a row of few (< 12) long, pointed setae; irregular submarginal row of blunt setae shorter than those on the dorsal margin; ventral margin with short, pointed setae and an irregular submarginal row of longer and blunt setae; central area of upper surface with very few short blunt setae (Fig. 50A). Fore tibia with several rows of long simple or feathered setae on ventral margin, dorsal margin with few hair-like setae; middle tibia with two rows of long, simple, pointed setae on ventral margin, dorsal margin with few hair-like setae; hind tibia with two rows of long and short feathered and simple setae on ventral margin, outer margin with short pointed setae and few hair-like setae. Tarsal claw straight with 2–4 denticles progressively larger apically (Fig. 50B).

Abdomen. Gills II–VII with dorsal and ventral lamellae, ventral lamella smaller. Gill I broken; gill II–VI with six or seven filaments on dorsal lamella and four on ventral lamella; gill VII with five filaments on dorsal lamella and three or four on ventral lamella. Posterior margin of terga IX with denticles. Posterior margin of terga VIII without or with minute denticles. Posterior margin of terga V–VII without denticles. Posterolateral projections present on segments VIII–IX.

This species is dedicated to Dr Marcela Miranda de Lima (Viçosa, Brazil).

Paratypes. • Papua New Guinea two nymphs in ethanol, GBIFCH01523467, one nymph on two slides GBIFCH01223153 (whole body), GBIFCH01223154 (gills), three female imagos in ethanol, GBIFCH01523468, Morobe Province, Bulolo River, east of Wau, 985 m, approx 7°20'16"S,146°43'42"E, 15–27.X.1964, W.L & J.G. Peters col (MZL).

Body length, male: 6.5–7 mm.

Medial emargination of labrum narrow with rounded denticles; ventral face of labrum with scattered thin setae; femora pale brown with two transverse dark brown bands in middle and at apex; posterior margin of abdominal segments V–VIII with well-developed denticles.

Head. Greyish brown, oval pale brown maculae in front of the median ocellus. Antenna yellowish, pedicel greyish brown. Compound eyes: lower portion black, upper portion orangish. Mouth parts yellowish brown. Thorax. Pronotum brownish orange, two triangular maculae on each side of the sagittal line (Fig. 51A). Mesonotum pale brown, washed with grey laterally. Femur greyish brown, apex pale brown, a pale brown band in the distal 1/3. Tibia yellowish, apex ringed with black. Tarsi yellowish. Abdomen. Terga medium brown, posterior margin of terga dark brown, as well as lateral sides. Sterna yellowish brown, posterior margin of sterna greyish brown laterally, nervous ganglia slightly tinted with greyish purple. Gills greyish purple. Terminal filaments pale brown, yellowish at apex.

Head. Labrum cordiform. Ratio labrum width/clypeus width 1.13. Ratio labrum width/insertion width 1.35. Medial emargination narrow with rounded denticles. Proximal row of setae present, simple. Number of setae on proximal row ~ 95–100. Distal row of setae simple. Median part of labrum in ventral view with scattered thin setae. Mandibles. Outer margin with row of setae on entire margin. Ending of the row near incisor. Right mandible with 7–9 setae below the mola. Maxillae. Posterior margin of cardo with numerous thin and stout setae. Proximo-lateral margin of stipes twisted. Apical-ventral row with 25–27 pectinate setae. Maxillary palp segment II 0.88–0.93 as long as segment I. Maxillary palp segment III 0.68–0.79 as long as segment I (Fig. 51B). Stout setae on outer margin of segment I present. Segment II inner margin without stout setae, outer margin without stout setae. Segment III inner margin without stout setae, dorsal surface fully covered with setae. Segment III 1.67–1.80 × as long as base width. Labium. Labial palp segment II 0.80–0.81 as long as segment I. Labial palp segment III 0.62–0.63 as long as segment I. Segment I inner margin with 17–20 stout setae, outer margin with 14–16 stout setae. Segment II outer margin with 17–20 stout setae. Segment III dorsal face with four stout setae. Segment III 2.97–3.02 × as long as base width. Hypopharynx. Apex of superlingua truncate.

Thorax. Dorsal margin of femur with a row of long, stout, pointed setae; irregular submarginal rows of shorter and pointed setae on fore femur, regular row of shorter, pointed setae on hind femur; ventral margin with short, stout setae and a submarginal row of long, pointed stout setae, except on hind leg where those of the submarginal row are short, pointed; central area of upper surface with long, pointed setae, shorter on hind femur. Fore tibia with several rows of long, simple setae on ventral margin, outer margin with few hair-like setae; middle tibia with two rows of long, simple pointed setae on ventral margin, outer margin with few hair-like setae; hind tibia with a row of simple long, thin setae on ventral margin, upper and lower surface covered with simple and feathered setae, outer margin with short and long pointed stout setae and hair-like setae. Tarsal claw hooked, with seven or eight denticles progressively larger apically (Fig. 51C).

Abdomen. Gills I–VII with dorsal and ventral lamellae, ventral lamella smaller. Gill I with ~ eight filaments on upper lamella and five on lower lamella, gills II–V with ~ 8–10 filaments on each lamella, gill VI with 5–7 filaments on each lamella, gill VII with four to five filaments on each lamella. Posterior margin of terga V–IX with denticles (Fig. 51D–G). Posterolateral projections present on segments VII–IX.

Eggs. General shape rounded. Size 140–145 μm × 85–90 μm (Fig. 51H). Chorionic surface smooth. No attachment structures visible. Micropyle not visible.

Holotype. • Indonesia, Papua, one nymph on slide, Sorong Island, 95 m, 19.II.2006, 00°49.354'S, 131°24.196'E, M. Balke & Tindige col [BH20], GBIFCH01223147 (MZB). Paratypes. • Indonesia, Papua, one nymph on slide, GBIFCH01223148, one nymph in ethanol, GBIFCH01523469 same data as holotype (MZL).

Body length, female: 4.5 mm.

Femora uniformly dark brown, without transverse bands; tarsal claw with only three teeth; all gills uni-lamellate.

Head. Vertex greyish brown, lighter between ocelli, frons pale brown washed with grey. Antenna yellowish, scape greyish. Mouth parts greyish brown. Thorax. Pronotum medium brown, washed with dark brown laterally, medially, and anteromedially. Mesonotum medium brown, dark brown laterally, with two blackish lines originating from the sides and reaching the posterior margin (Fig. 36C). Femur greyish brown, lighter at base. Tibia yellowish, fore tibia washed with grey apically. Tarsi yellowish. Abdomen. Terga medium brown, a lighter sagittal line increasing in width posteriorly. Sterna yellowish, segments I–VII washed with greyish brown laterally. Gills greyish brown. Terminal filaments yellowish.

Head. Labrum cordiform (Fig. 52A). Ratio labrum width/clypeus width 1.02. Ratio labrum width/insertion width 1.45. Medial emargination shallow without denticles. Proximal row of setae present, simple. Number of setae on proximal row ~ 30–40. Distal row of setae present, simple. Median part of labrum in ventral view with scattered thin setae (Fig. 52B). Mandibles. Outer margin with row of setae on distal 1/2. Ending of the row before incisor. Right mandible with six setae below the mola (Fig. 52C). Maxillae. Posterior margin of cardo broken. Proximo-lateral margin of stipes with one medium size and two short stout setae (Fig. 34I). Apical-ventral row with 20–25 pectinate setae. Maxillary palp segment II 0.76–1.0 as long as segment I. Maxillary palp segment III 0.76–0.9 as long as segment I (Fig. 52D). Stout setae on outer margin of segment I absent. Segment II inner margin with two or three stout setae. Segment II outer margin with two stout setae. Segment III inner margin with three or four stout setae, dorsal surface with few setae at apex. Segment III 1.86–2.07 × as long as base width. Labium. Labial palp segment II 0.73–0.75 as long as segment I. Labial palp segment III 0.67–0.80 as long as segment I (Fig. 52E). Segment I inner margin with 7–9 stout setae, outer margin with 6–9 stout setae. Segment II, outer margin with 2–4 stout setae. Segment III, dorsal face with two or three stout setae. Segment III 2.88–3.33 × as long as base width. Hypopharynx. Apex of superlingua truncate (Fig. 52F).

Thorax. Dorsal margin of femur with a row of few (< 12) long and pointed setae; irregular submarginal row of blunt, pointed setae shorter or as long as those on the dorsal margin; ventral margin with short, pointed setae and an irregular submarginal row of longer, blunt setae; central area of upper surface with very few short pointed and blunt setae (Fig. 53A–B). Fore tibia with several rows of long simple or feathered setae on ventral margin, dorsal margin with numerous hair-like setae; middle tibia with one row of short, simple, pointed setae, and an irregular submarginal row of short, pointed setae on ventral margin, dorsal margin with numerous hair-like setae; hind tibia with two rows of long and short feathered and simple setae on ventral margin, dorsal margin with long pointed setae and numerous hair-like setae. Tarsal claw slightly hooked with three denticles progressively larger apically (Fig. 53C).

Abdomen. All gills uni-lamellate, ventral lamella lacking; gill I with two filaments (Fig. 53D), gills II–VI with four or five filaments (Fig. 53E), gill VII with two filaments (Fig. 53F). Posterior margin of terga IX with minute denticles. Posterior margin of terga V–VIII without denticles. Posterolateral projections present on segments VIII–IX.

The species name is to recognise the Moi people, a native tribe of Sorong, who fight to keep their intact environment and their way of living.

Holotype. • Papua New Guinea, one nymph on slide, Madang Province, Aiome area 130 m, 11.III.2007, 05°10.593'S, 144°42.800'E, Kinibel col [PNG156], GBIFCH01223146 (ZSM). Paratype. • Papua New Guinea, one nymph in ethanol, GBIFCH01523470, same data as holotype (MZL).

Body length, female: 5 mm.

Median emargination of labrum shallow with flat denticles; ventral surface of labrum with stout setae in row; femora pale brown with two transverse dark brown bands in middle and at apex; posterolateral expansions present on segments VII–IX.

Head. Medium brown, washed with greyish brown between ocelli, pale brown between lateral ocelli and compound eyes, occiput with pale brown vermiform maculae around the sagittal line, clypeus dark brown. Antenna broken. Mouth parts pale brown. Thorax. Pronotum pale brown, dark brown anteriorly and laterally, as well as along the sagittal line. Mesonotum pale brown, washed with dark brown laterally and along the distal half of the sagittal line, fore wing pads medium brown, with a dark brown spot at the base. Femur pale brown, with a large medium brown band in the middle and at apex. Tibia yellowish, with a dark brown ring at apex. Tarsi yellowish to pale brown. Abdomen. Terga segments I and II entirely dark brown, segments III and IV medium brown with a dark brown macula in anteromedian position, this macula with a light brown longitudinal line on segments V–VII, segments VII–X dark brown. Sterna yellowish grey, segments III–VI with a dark brown macula in anteromedian position. Gills pale grey, tinted with purple. Terminal filaments broken.

Head. Labrum cordiform (Fig. 54A). Ratio labrum width/clypeus width 0.95. Medial emargination shallow with flat denticles. Proximal row of setae present, simple. Number of setae on proximal row ~ 22. Distal row of setae present, simple. Median part of labrum in ventral view with stout setae in row (Fig. 54B). Mandibles. Outer margin with row of setae on distal 1/2. Ending of the row near incisor. Right mandible with eight setae below the mola (Fig. 54C). Maxillae. Posterior margin of cardo broken. Proximo-lateral margin of stipes with one long and stout seta. Apical-ventral row with 16 pectinate setae. Maxillary palp segment II 0.84 as long as segment I. Maxillary palp segment III 0.67 as long as segment I (Fig. 54D). Stout setae on outer margin of segment I absent. Segment II inner margin with four stout setae, outer margin with four stout setae. Segment III inner margin with four stout setae, dorsal surface with few setae at apex. Segment III 1.64 × as long as base width. Labium. Labial palp segment II 0.94 as long as segment I. Labial palp segment III 0.82 as long as segment I (Fig. 54E). Segment I inner margin with 14 stout setae, outer margin with 13 stout setae. Segment II outer margin with seven stout setae. Segment III dorsal face with six stout setae. Segment III 3.11 × as long as base width. Hypopharynx. Apex of superlingua truncate (Fig. 54F).

Thorax. Dorsal margin of fore femur with long, stout, pointed setae, and a submarginal row of long, stout, pointed setae; dorsal margin of mid- and hind femur with a row of long, stout, pointed setae; a submarginal row of pointed setae as long as or a little bit shorter than those on the dorsal margin; ventral margin with short, stout setae and an irregular submarginal row of long, pointed stout setae, except on hind leg where those on ventral margin are shorter; central area of upper surface with few short or long stout and pointed setae (Fig. 55A–B). Fore tibia with two or three rows of long simple or feathered setae on ventral margin, dorsal margin with numerous hair-like setae; middle tibia with two rows of long, simple, pointed setae on ventral margin, outer margin with numerous hair-like setae; hind tibia with two rows of simple and feathered long, stout setae on ventral margin, outer margin with a row of long and short simple stout setae, with numerous hair-like setae. Tarsal claw hooked with 4–6 denticles progressively larger apically (Fig. 55C).

Abdomen. Gill I with both lamellae. Gills II–VII with dorsal and ventral lamellae of the same size. Gills very small, gill I with five or six filaments on each lamella, gills II–VI with 7–10 filaments on each lamella (Fig. 55D). Posterior margin of terga IX with denticles. Posterior margin of terga VIII without or with minute denticles. Posterior margin of terga V–VII without denticles. Posterolateral projections present on segments VII–IX.

The species name is a Latin word composed of silva (forest) and pumilo (dwarf) to recognise the Melanesian Dwarf tribe of Aiome, and the land where the specimens of this new species come from.

• Indonesia, Papua, one nymph on slide, GBIFCH01223145, Road Nabire–Enarotali KM 55, 774 m, 22.X.2011, 03°29.796'S, 135°43.885'E, M. Balke col [PAP9] (MZL).

Nymph. Labrum trapezoidal; dorsal face of labrum with simple row in distal position; femora pale brown with two transverse dark brown bands in middle and at apex.

Femur yellowish brown, with a large median greyish brown stripe and another one at apex (Fig. 57A–B). Tibia yellowish. Tarsi yellowish.

Head. Labrum trapezoid (Fig. 56A). Ratio labrum width/clypeus width 1.11. Ratio labrum width/insertion width 1.56. Medial emargination narrow with flat denticles. Proximal row of setae present, simple. Number of setae on proximal row ~ 46. Distal row of setae present, simple. Median part of labrum in ventral view with scattered thin setae (Fig. 56B). Mandibles. Outer margin with row of setae on distal 1/2. Ending of the row before incisor. Right mandible with seven setae below the mola (Fig. 56C). Maxillae. Posterior margin of cardo with few hair-like setae, together with one stout and long seta in submarginal position. Proximo-lateral margin of stipes with a bunch of ~ six stout setae increasing in size distally (Fig. 34H). Apical-ventral row with 32 pectinate setae. Maxillary palp segment II 0.73–0.78 as long as segment I. Maxillary palp segment III 0.79–0.81 as long as segment I (Fig. 56D). Stout setae on outer margin of segment I absent. Segment II inner margin with four stout setae, outer margin with four stout setae. Segment III inner margin with six stout setae, dorsal surface with few setae at apex. Segment III 2.36 × as long as base width. Labium. Labial palp segment II 0.79 as long as segment I. Labial palp segment III 0.67 as long as segment I (Fig. 56E). Segment I inner margin with nine stout setae, outer margin with ten stout setae. Segment II outer margin with five stout setae. Segment III dorsal face with three stout setae. Segment III 3.00 × as long as base width. Hypopharynx. Apex of superlingua truncate (Fig. 56F).

Thorax. All legs much longer than in any other species of the Thraulus lineage; fore femur 3 × longer than wide (Fig. 57A), hind femur 4 × longer than wide (Fig. 57B). Dorsal margin of femur with a row of long, pointed setae; regular submarginal row of blunt setae shorter than those on the dorsal margin; ventral margin with short, pointed setae and an irregular submarginal row of longer, pointed setae; central area of upper surface with few long and short blunt setae. Fore tibia with several rows of long simple or feathered setae on ventral margin, dorsal margin with few hair-like setae; middle tibia with two rows of short, simple, pointed setae on ventral margin, dorsal margin with numerous hair-like setae; hind tibia with two or three rows of long and short feathered and simple setae on ventral margin, outer margin with short, pointed setae and numerous hair-like setae. Tarsal claw hooked with four or five denticles progressively larger apically (Fig. 57C–D).

Abdomen. Most gills missing. Gill IV very long, uni-lamellate, with 5–6 filaments (Fig. 57E). Filamentous projections on apex. Posterior margin of terga IX with minute denticles. Posterior margin of terga V–VIII without denticles. Posterolateral projections present on segments VIII–IX.

• Papua New Guinea, one nymph on slide, GBIFCH01223128, Western Highlands Province, Simbai, 1800–2000 m, 25.II.2007, 05°16.330'S, 144°33.176'E, Kinibel col [PNG133] (MZL).

Nymph. Labrum trapezoidal; dorsal face of labrum with multiple rows in distal position; femora pale brown with two transverse dark brown bands in middle and at apex.

Femur yellowish, a medium brown transversal band in the middle, apex medium brown (Fig. 59A–B). Tibia yellowish, apex dark brown. Tarsi yellowish. Gills greyish brown.

Head. Labrum trapezoid (Fig. 58A). Ratio labrum width/clypeus width 1.17. Ratio labrum width/insertion width 1.46. Medial emargination narrow with flat denticles. Proximal row of setae present, simple. Number of setae on proximal row ~ 106. Distal row of setae multiple. Median part of labrum in ventral view with scattered thin setae (Fig. 58B). Mandibles. Outer margin with row of setae on distal 1/2. Ending of the row near incisor. Right mandible with nine setae below the mola (Fig. 58C). Maxillae. Posterior margin of cardo broken. Proximo-lateral margin of stipes with a bunch of ~ ten stout short and medium size setae. Apical-ventral row with 30 pectinate setae. Maxillary palp segment II 0.76 as long as segment I. Maxillary palp segment III 0.71 as long as segment I (Fig. 58D). Stout setae on outer margin of segment I absent. Segment II inner margin with two stout setae, outer margin without stout setae. Segment III inner margin with one stout seta, dorsal surface half covered with setae. Segment III 1.50 × as long as base width. Labium. Labial palp segment II 0.79 as long as segment I. Labial palp segment III 0.65 as long as segment I (Fig. 58E). Segment I inner margin with 14 stout setae, outer margin with 15 stout setae. Segment II outer margin with nine stout setae. Segment III dorsal face with two stout setae. Segment III 2.73 × as long as base width. Hypopharynx. Apex of superlingua truncate (Fig. 58F).

Thorax. Dorsal margin of femur with a row of very long, thin, pointed setae; regular submarginal row of pointed setae as long as or shorter than those on the dorsal margin, multiple submarginal rows on fore femur (Fig. 59A); ventral margin with short, pointed setae and an irregular submarginal row of long, pointed setae, except on hind femur where they are short and stout; central area of upper surface with very long pointed setae on fore femur, long and short on middle femur and short on hind femur (Fig. 59B). Fore tibia with two or three rows of long simple or feathered setae on ventral margin, dorsal margin with numerous hair-like setae; middle tibia with a row of short, simple, pointed setae on ventral margin and a submarginal row of long, simple setae, outer margin with numerous hair-like setae; hind tibia with a marginal row of simple short, thin setae and a submarginal row of long to very long simple setae on ventral margin, outer margin with long to very long pointed setae and numerous hair-like setae. Tarsal claw slightly hooked, with five or six denticles progressively larger, except distal much larger (Fig. 59C).

Abdomen. Gill I with both lamellae. Gills II–VII with dorsal and ventral lamellae, ventral lamella smaller. Gill I with nine or ten filaments on dorsal lamella and five or six on ventral lamella, gills II–VI with 11–12 filaments on dorsal lamella and six or seven on ventral lamella (Fig. 59E), gill VII with nine or ten filaments on dorsal lamella and four or five on ventral lamella. Posterior margin of terga IX with denticles (Fig. 59D). Posterior margin of terga V–VIII without denticles. Posterolateral projections present on segments VIII–IX.

Gill I similar in shape to the following ones. Ventral lamella of gills smaller but not highly reduced, with ≥ 9 processes; claws with 6–10 teeth; posterior margin of abdominal segment VIII with small denticles; posterolateral expansions on abdominal segments VII–IX, VI–IX or even V–IX. Eggs fusiform with hook-like attachment structures at each pole.

From the greek κόσμημα meaning jewel and νύμφη meaning nymph, to recognise the peculiar beauty of the species in this genus. The gender is feminine.

Kosminympha sabrinae sp. nov. by present designation.

Kosminympha asarorum sp. nov., K. balkei sp. nov., K. baruya sp. nov., K. kalamorum sp. nov., K. paulinae sp. nov., K. sabrinae sp. nov.

Size: 10.0–11.0 mm for male mature nymphs; 10.5–12.5 mm for mature female nymphs.

Head. Prognathous; labrum cordiform or trapezoid, anterior emargination either shallow with flat denticles or narrow with pointed denticles; proximal row of setae long, with 80–140 setae; distal row of setae multiple; median part of labrum in ventral view with scattered thin and long setae or with stout and short setae in a row. Outer margin of mandibles with a row of setae. Proximo-lateral margin of stipes with a bunch of setae, either long and thin or short and stout. Apex of superlingua of hypopharynx either truncate or emarginate. Thorax. Central area of upper face of femora with numerous short or long, pointed, or blunt setae; tarsal claw hooked, with 6–10 teeth. Abdomen. Gill I similar in shape to the following ones; gill I composed of dorsal and ventral lamellae, each with 9–15 filaments; gills II–VI composed of dorsal and ventral lamellae, each with 15–25 filaments; gill VII composed of dorsal and ventral lamellae, each with 5–15 filaments, depending on species. Posterior margin of abdominal terga II–VIII without or with minute denticles; posterolateral expansions of the abdomen present on segments V–IX, VI–IX or VII–IX. Egg. Elongated, with attachments structures fixed at each pole, composed of a long branch ending by a double fishhook.

Among mayfly nymphs of the subfamily Choroterpinae and of the tribe Thraulini, the genus Kosminympha is most similar to Nonnullidens by its gill composition. It differs anyway by the less reduced gill ventral lamella, bearing ≥ nine filaments whereas in Nonnullidens, ventral lamella bears at most eight filaments (in N. mariae, it can also reach ten filaments). Posterolateral projections of the abdomen are present on segments V–IX, VI–IX or VII–IX, whereas they are present on segments VIII–IX in Nonnullidens (VII–IX in some species). Posterior margin of tergite VIII always bears small denticles, whereas they are absent in Nonnullidens (present and well-developed in N. mariae). Finally, the eggs of Kosminympha are similar in shape to those of the genus Thraulus, being elongated and bearing attachment structures at each pole, whereas they are rounded and without attachment structure in Nonnullidens.

The genus Kosminympha is only known from Papua New Guinea and has been found on the eastern part of the island; the genus has yet not been recorded from Indonesia Papua.

Holotype. • Papua New Guinea, one nymph on slide, Eastern Highlands Province, Akameku – Brahmin, Bismarck Range, 2200 m, 23.XI.2006, 05°56.801'S, 145°22.238'E, M. Balke & Kinibel col [PNG106], GBIFCH01223156 (ZSM). Paratype. • Papua New Guinea, one nymph in ethanol, GBIFCH01523450, same data as holotype (MZL).

Body length, female: 10.5 mm.

Labrum trapezoid; dorsal surface of labrum with distal row multiple; tarsal claw hooked with seven teeth; abdominal tergites predominantly medium to dark brown; central area of upper surface of femora with numerous short, stout, and blunt setae, shorter on hind femur; posterior margin of abdominal segment VII without denticles.

Head. Greyish brown, pale brown between lateral ocelli and compound eyes, pale brown in front of the fore ocellus, clypeus medium brown. Antenna broken. Mouth parts greyish brown. Thorax. Pronotum pale brown, dark brown laterally and anteriorly, with a double triangular macula along the sagittal line. Mesonotum pale brown, washed with greyish brown, fore wing pads pale brown (Fig. 60C). Femur greyish brown, whitish at base and apex. Tibia uniformly yellowish brown. Tarsi yellowish brown. Abdomen. Terga greyish brown, posterior margin and sagittal line pale brown. Sterna yellowish. Gills upper lamella blackish, lower lamella greyish. Terminal filaments broken.

Head. Labrum trapezoid (Fig. 62A). Ratio labrum width/clypeus width 1.25. Ratio labrum width/insertion width 1.64. Medial emargination shallow with flat denticles. Proximal row of setae present, shifted to distal margin. Number of setae on proximal row ~ 115. Distal row of setae multiple. Median part of labrum in ventral view with scattered thin setae (Fig. 62B). Mandibles. Outer margin with row of setae on distal 1/2. Ending of the row before incisor. Right mandible with 11 setae below the mola (Fig. 62C). Maxillae. Posterior margin of cardo with few stout and medium size setae together with some hair-like setae and two or three stout and long setae in submarginal position. Proximo-lateral margin of stipes with a bunch of ~ 15 medium size and stout setae. Apical-ventral row with 36 pectinate setae. Maxillary palp segment II 0.75 as long as segment I. Maxillary palp segment III 0.63 as long as segment I (Fig. 62D). Stout setae on outer margin of segment I absent. Segment II inner margin with four stout setae, outer margin without stout setae. Segment III inner margin with four stout setae, dorsal surface with few setae at apex. Segment III 1.73 × as long as base width. Labium. Labial palp segment II 0.92 as long as segment I. Labial palp segment III 0.65 as long as segment I (Fig. 62E). Segment I inner margin with 11 stout setae, outer margin with 18 stout setae. Segment II outer margin with ten stout setae. Segment III dorsal face with five stout setae. Segment III 2.77 × as long as base width. Hypopharynx. Apex of superlingua truncate (Fig. 62F).

Thorax. Dorsal margin of femur with a row of very long, stout, pointed setae (Fig. 63A); an irregular submarginal row of pointed setae as long as or a little bit shorter than those on the dorsal margin, more regular on hind femur; ventral margin with short, stout setae and an irregular submarginal row of long, pointed stout setae, except on hind leg where the submarginal row is regular (Fig. 63B); central area of upper surface with numerous short, stout, blunt setae. Fore tibia with two or three rows of long simple or feathered setae on ventral margin, dorsal margin with numerous hair-like setae; middle tibia with a row of short, simple, pointed setae on ventral margin and a submarginal row of long, simple setae, outer margin with numerous hair-like setae; hind tibia with two or three rows of simple long, stout setae on ventral margin, lower surface with few simple, thin setae, outer margin with short and long pointed stout setae and hair-like setae. Tarsal claw hooked with seven denticles progressively larger apically, except distal much smaller (Fig. 63C).

Abdomen. Gill I with both lamellae. Gills II to VII with dorsal and ventral lamellae of the same size. Gill I with ~ 12 filaments on each lamella, gills II–VI with ~ 20 filaments on each lamella (Fig. 63D), gill VII with ~ ten filaments on each lamella. Filamentous projections on apex. Posterior margin of terga IX with denticles. Posterior margin of terga VIII with minute denticles. Posterior margin of terga V–VII without denticles. Posterolateral projections present on segments VI–IX.

The specific name is to acknowledge the Asaro tribe or “Mudmen” and their living traditions, who inhabit close to where the specimens were collected.

Holotype. • Papua New Guinea, one nymph on slide, Eastern Highlands Province, Marawaka, Ande, 1700–1800 m, 9.XI.2006, approx. 07°01.697'S, 145°49.807'E, M. Balke & Kinibel col [PNG87], GBIFCH01223158 (ZSM). Paratypes. • Papua New Guinea, one nymph on slide GBIFCH01223157, 2 nymphs in ethanol, GBIFCH01523449, same data as holotype (MZL).

Body length, female: 10.5 mm.

Labrum cordiform; median part of labrum in ventral view with a row of stout and short setae; abdominal tergites predominantly orange with dark lateral marks; gills II–VI with ~ 15 filaments on each lamella; posterior margin of abdominal segment VII with minute denticles; posterolateral expansions present on abdominal segments V–IX.

Head. Medium brown, washed with greyish brown between ocelli, pale brown between lateral ocelli and compound eyes, clypeus medium brown. Antenna uniformly yellowish. Mouth parts brownish. Thorax. Pronotum medium brown, pale brown laterally, two dark brown arch-like maculae close to the sagittal line. Mesonotum pale brown, medium brown on antero-lateral corners, fore wing pads medium brown (Fig. 60A). Femur uniformly yellowish brown, fore femur with proximal 1/3 whitish. Tibia yellowish orange. Tarsi yellowish orange. Abdomen. Terga brownish orange, segment II–VII with a pair of more or less triangular dark brown maculae along the sagittal line, segment IX with a pair of square dark brown maculae, segment X entirely brownish orange. Sterna yellowish from segments I–IV, becoming brownish yellow to brownish orange posteriorly. Gills purple. Terminal filaments yellowish, darker near base.

Head. Labrum cordiform (Fig. 65A). Ratio labrum width/clypeus width 0.96. Ratio labrum width/insertion width 1.24–1.30. Medial emargination narrow with pointed denticles. Proximal row of setae present, simple. Number of setae on proximal row ~ 82–93. Distal row of setae multiple. Median part of labrum in ventral view with stout setae in row (Fig. 65B). Mandibles. Outer margin with row of setae on distal 1/2. Ending of the row near incisor. Right mandible with 15–22 setae below the mola (Fig. 65C). Maxillae. Posterior margin of cardo with many hair-like and long setae, together with two or three long and stout setae in submarginal position. Proximo-lateral margin of stipes with a bunch of ten long hair-like setae (Fig. 34K). Apical-ventral row with 14–15 pectinate setae. Maxillary palp segment II 0.80–0.81 as long as segment I. Maxillary palp segment III 0.57–0.65 as long as segment I (Fig. 65D). Stout setae on outer margin of segment I absent. Segment II inner margin with 1–3 stout setae, outer margin without stout setae. Segment III inner margin with four stout setae, dorsal surface half covered with setae. Segment III 1.53–1.56 × as long as base width. Labium. Labial palp segment II 0.88 as long as segment I. Labial palp segment III 0.76–0.80 as long as segment I (Fig. 65E). Segment I inner margin with 25–35 stout setae, outer margin with 28–30 stout setae. Segment II outer margin with 12 stout setae. Segment III dorsal face with six or seven stout setae. Segment III 2.53–2.71 × as long as base width. Hypopharynx. Apex of superlingua emarginated (Fig. 65F).

Thorax. Dorsal margin of femur with a row of very long, thin, pointed setae; irregular submarginal rows of pointed setae as long as or a little bit shorter than those on the dorsal margin, these setae decrease in size from the fore- to the hind femur (Fig. 66A–B); ventral margin with short, stout setae and irregular submarginal rows of long, pointed stout setae, except on hind leg where those on ventral margin are shorter; central area of upper surface with numerous long, stout, pointed setae, shorter on hind femur. Fore tibia with several rows of long simple or feathered setae on ventral margin, dorsal margin with numerous hair-like setae; middle tibia with two rows of long, simple, pointed setae on ventral margin, outer margin with numerous hair-like setae; hind tibia with several rows of simple and feathered long, stout setae on ventral margin, lower surface covered with simple and feathered setae, outer margin with short and long pointed stout setae and hair-like setae. Tarsal claw hooked with eight or nine denticles progressively larger apically (Fig. 66C).

Abdomen. Gill I with both lamellae. Gills II–VII with dorsal and ventral lamellae of the same size. Gill I with ~ 10 filaments on each lamella, gills II–VI with ~ 15 filaments on each lamella, gill VII minute with 5–7 filaments on each lamella. Filamentous projections on apex. Posterior margin of terga IX with denticles. Posterior margin of terga VII–VIII with minute denticles. Posterior margin of terga V–VI without denticles. Posterolateral projections present on segments V–IX.

This species is dedicated to Dr Michael Balke (ZSM, Germany), collector of most of the species treated here.

Holotype. • Papua New Guinea, nymph on slide, Central Province, Kokoda trek, 1400 m, I.2008, 09°01.952'S, 147°44.455'E, Posman col [PNG172], GBIFCH01223159 (ZSM). Paratypes. • Papua New Guinea, one nymph in ethanol, GBIFCH01523446, same data as holotype (MZL). • Papua New Guinea, one nymph on slide, GBIFCH01223160, 2 nymphs in ethanol, GBIFCH01523445, Gulf Province, Marawaka, Mala, 1400 m, 11.XI.2006, 07°05.664'S, 145°44.467'E, M. Balke and Kinibel col [PNG90] (MZL).

Body length, female: 8.5 mm.

Labrum cordiform; emargination narrow with pointed denticles; tarsal claw straight with nine or ten teeth; abdominal tergites predominantly medium to dark brown; posterolateral expansions present on abdominal segments VII–IX.

Head. Greyish brown, pale brown between antennae and front ocellus. Antenna broken. Mouth parts pale brown, labrum and mandibles dark brown. Thorax. Pronotum greyish brown, dark brown laterally and medially. Mesonotum pale brown, with greyish maculae as in Fig. 61A. Femur greyish brown, yellowish proximally and subdistally. Tibia uniformly yellowish. Tarsi uniformly yellowish. Abdomen. Terga uniformly medium brown, with pale brown dots close to gills insertion. Sterna yellowish in median part, posterior margin greyish laterally. Gills brownish purple. Terminal filaments yellowish, darker near base.

Head. Labrum cordiform (Fig. 67A). Ratio labrum width/clypeus width 1.06–1.08. Ratio labrum width/insertion width 1.28. Medial emargination narrow with pointed denticles. Proximal row of setae present, simple. Number of setae on proximal row ~ 120–140. Distal row of setae multiple. Median part of labrum in ventral view with scattered thin setae (Fig. 67B). Mandibles. Outer margin with row of setae on entire margin. Ending of the row near incisor. Right mandible with 14–16 setae below the mola (Fig. 67C). Maxillae. Posterior margin of cardo with many hair-like setae. Proximo-lateral margin of stipes with a bunch of ~ 15 stout, long, pointed setae (Fig. 34L). Apical-ventral row with 22–24 pectinate setae. Maxillary palp segment II 0.82–0.88 as long as segment I. Maxillary palp segment III 0.75–0.85 as long as segment I (Fig. 67D). Stout setae on outer margin of segment I present. Segment II inner margin with 1–5 stout setae, outer margin with 2–7 stout setae. Segment III inner margin with four or five stout setae, dorsal surface fully covered with setae. Segment III 1.67–1.77 × as long as base width. Labium. Labial palp segment II 0.77–0.83 as long as segment I. Labial palp segment III 0.60–0.62 as long as segment I (Fig. 67E). Segment I inner margin with 23–25 stout setae, outer margin with 33–38 stout setae. Segment II outer margin with 21–22 stout setae. Segment III dorsal face with three or four stout setae. Segment III 2.56–2.67 × as long as base width. Hypopharynx. Apex of superlingua emarginated (Fig. 67F).

Thorax. Dorsal margin of femur with a row of very long, thin, pointed setae; irregular submarginal rows of pointed setae as long as or a little bit shorter than those on the dorsal margin, ventral margin with short, stout setae and submarginal rows of long, pointed stout setae, except on hind leg where those on ventral margin are shorter; central area of upper surface with numerous long, stout, pointed setae, shorter on hind femur (Fig. 68A–C). Fore tibia with several rows of long simple or feathered setae on ventral margin, dorsal margin with numerous hair-like setae; middle tibia with several rows of long, simple, pointed setae on ventral margin, outer margin with numerous hair-like setae; hind tibia with several rows of simple and feathered long, stout setae on ventral margin, lower surface covered with simple and feathered setae, outer margin with short and long pointed stout setae and hair-like setae. Tarsal claw hooked with nine or ten denticles progressively larger, except distal much smaller (Fig. 68D).

Abdomen. Gill I with both lamellae. Gills II–VII with dorsal and ventral lamellae of the same size. Gill I with nine or ten filaments on each lamella (Fig. 68J), gills II–VI with ~ 15 filaments on each lamella (Fig. 68L), gill VII with three or four filaments on each lamella (Fig. 68K). Filamentous projections on apex. Posterior margin of terga VII–IX with denticles (Fig. 68G–I). Posterior margin of terga V–VI with minute denticles (Fig. 68E–F). Posterolateral projections present on segment VII–IX.

The specific name is to recognise the tribe Baruya living in the valleys of Wonemara and Marawaka, where some of the specimens come from, and is a noun in apposition.

Holotype. • Papua New Guinea, one nymph on slide, Western Highlands Province, Simbai, 1800–2000 m, 26.II.2007, 05°15.872'S, 144°32.717'E, Kinibel col [PNG134], GBIFCH01223161 (ZSM). Paratypes. • Papua New Guinea, one nymph on slide, GBICH01223162, 2 nymphs in ethanol, GBIFCH01523447, same data as holotype (MZL). • Papua New Guinea, one nymph on slide, GBIFCH01223163, Eastern Highlands Province, Akameku – Brahmin, Bismarck Range, 2200 m, 23.XI.2006, 05°56.801'S, 145°22.238'E, M. Balke & Kinibel col [PNG106] (MZL).

Body length, female: 11.5 mm.

Labrum trapezoid; median part of labrum in ventral view with scattered thin and long setae; abdominal tergites predominantly orange with dark lateral marks; posterior margin of abdominal segment VII without denticles; posterolateral expansions present on abdominal segments VII–IX.

Head. Medium brown, washed with dark brown between ocelli, pale brown between lateral ocelli and compound eyes, clypeus medium brown. Antenna uniformly yellowish. Mouth parts brownish. Thorax. Pronotum medium brown, dark brown laterally, a large dark brown anchor-like macula straddling the sagittal line. Mesonotum pale brown, medium brown laterally, fore wing pads dark brown (Fig. 60B). Femur: fore femur yellowish brown, with proximal 1/3 whitish, mid- and hind femur pale brown. Tibia yellowish brown, apex of fore tibia dark brown. Tarsi pale brown. Abdomen. Terga brownish orange, segment II–V with a pair of elongated dark brown maculae along the sagittal line, segments VI–IX with a pair of triangular to square dark brown maculae, segment IX with two small dark stripes along the lateral margins, segment X entirely brownish orange. Sterna yellowish from segments I–IV, becoming brownish yellow to brownish orange posteriorly. Gills purple. Terminal filaments yellowish.

Head. Labrum trapezoid (Fig. 69A). Ratio labrum width/clypeus width 1.26–1.27. Ratio labrum width/insertion width 1.69–1.71. Medial emargination shallow with flat denticles. Proximal row of setae simple, shifted to distal margin. Number of setae on proximal row ~ 123–130. Distal row of setae multiple. Median part of labrum in ventral view with scattered thin setae (Fig. 69B). Mandibles. Outer margin with row of setae on distal 1/2. Ending of the row before incisor. Right mandible with 8–13 setae below the mola (Fig. 69C). Maxillae. Posterior margin of cardo with few stout and medium size setae (Fig. 34D). Proximo-lateral margin of stipes with a bunch of eight long and stout setae. Apical-ventral row with 34–44 pectinate setae. Maxillary palp segment II 0.77–0.86 as long as segment I. Maxillary palp segment III 0.77–0.86 as long as segment I (Fig. 69D). Stout setae on outer margin of segment I absent. Segment II inner margin with two stout setae, outer margin without or with up to two stout setae. Segment III inner margin with six stout setae, dorsal surface with few setae at apex. Segment III 1.67–2.00 × as long as base width. Labium. Labial palp segment II 0.84–0.87 as long as segment I. Labial palp segment III 0.60–0.64 as long as segment I (Fig. 69E). Segment I inner margin with 8–10 stout setae. Segment I outer margin with 12–16 stout setae. Segment II outer margin with 6–10 stout setae. Segment III dorsal face without stout setae. Segment III 2.56–3.33 × as long as base width. Hypopharynx. Apex of superlingua truncate (Fig. 69F).

Thorax. Dorsal margin of fore femur with few stout, short, pointed setae, and a double submarginal row of long, stout, pointed setae (Fig. 70A); dorsal margin of mid- and hind femur with a row of long, thin, pointed setae; a submarginal row of pointed setae as long as or a little bit shorter than those on the dorsal margin (Fig. 70B); ventral margin with short, stout setae and an irregular submarginal row of long, pointed stout setae, except on hind leg where those on ventral margin are shorter; central area of upper surface with numerous short, stout, blunt setae. Fore tibia with two or three rows of long simple or feathered setae on ventral margin, dorsal margin with numerous hair-like setae; middle tibia with one row of short, simple, pointed setae, and a submarginal row of long, simple, pointed stout setae on ventral margin, outer margin with numerous hair-like setae; hind tibia with a row of simple long, stout setae on ventral margin, outer margin with short pointed stout setae and hair-like setae. Tarsal claw hooked with 6–8 denticles progressively larger apically, followed by two tiny protuberances (Fig. 70C).

Abdomen. Gill I with both lamellae. Gills II–VII with dorsal and ventral lamellae of the same size. Gill I with ~ 10 filaments on each lamella, gills II–VI with ~ 15 filaments on each lamella (Fig. 70F), gill VII minute with 5–7 filaments on each lamella. Filamentous projections on apex. Posterior margin of terga IX with minute denticles, or with larger denticles (Fig. 70D). Posterior margin of terga VIII with minute denticles (Fig. 70E). Posterior margin of terga V–VII without denticles. Posterolateral projections present on segments VII–IX.

The specific name is to acknowledge the Kalam tribe of the remote area of Simbai, where the specimens described here come from.

Holotype. • Papua New Guinea, nymph on slide, Morobe Province, Menyamya, Mt Inji, 1700 m, 14.XI.2006, approx. 07°14.813'S, 146°01.330'E, M. Balke & Kinibel col [PNG96], GBIFCH01223164 (ZSM). Paratype. • Papua New Guinea, one nymph in ethanol, GBIFCH01523448, same data as holotype (MZL).

Body length, female: 8.2 mm.

Labrum trapezoid; dorsal surface of labrum with distal row simple; tarsal claw hooked with seven teeth; abdominal tergites predominantly medium to dark brown; central area of upper surface of femora with numerous long, stout, and blunt setae, shorter on hind femur; posterior margin of abdominal segment VII with minute denticles.

Head. Greyish brown, pale brown in front of the fore ocellus, whitish between lateral ocelli and compound eyes. Antenna whitish, scape and pedicel greyish white. Mouth parts greyish. Thorax. Pronotum medium brown, dark brown laterally and anteriorly as in Fig. 61C. Mesonotum medium brown bordered with dark brown, fore wing pads dark brown. Femur medium brown, except proximal part whitish. Tibia uniformly yellowish to pale brown. Tarsi yellowish to pale brown. Abdomen. Terga uniformly dark brown, sagittal line lighter. Sterna yellowish, segments VIII–IX darker. Gills greyish purple. Terminal filaments yellowish brown.

Head. Labrum trapezoid (Fig. 71A). Ratio labrum width/clypeus width 1.28. Ratio labrum width/insertion width 1.71. Medial emargination shallow with flat denticles. Proximal row of setae simple, shifted to anterior margin. Number of setae on proximal row ~ 120. Distal row of setae multiple, shifted to anterior margin. Median part of labrum in ventral view with scattered thin setae (Fig. 71B). Mandibles. Outer margin with row of setae on distal 1/2. Ending of the row before incisor. Right mandible with 5 setae below the mola (Fig. 71C). Maxillae. Posterior margin of cardo with ~ 10 stout pointed and long setae. Proximo-lateral margin of stipes with a bunch of ~ 12 medium size and short stout setae (Fig. 34J). Apical-ventral row with 45 pectinate setae. Maxillary palp segment II 0.86 as long as segment I. Maxillary palp segment III 0.67 as long as segment I (Fig. 71D). Stout setae on outer margin of segment I absent. Segment II inner margin without stout setae. Segment II outer margin without stout setae. Segment III inner margin with six stout setae, dorsal surface with few setae at apex. Segment III 1.87 × as long as base width. Labium. Labial palp segment II 0.75 as long as segment I. Labial palp segment III 0.59 as long as segment I (Fig. 71E). Segment I inner margin with ten stout setae. Segment I outer margin with 17 stout setae. Segment II outer margin with ten stout setae. Segment III dorsal face with two stout setae. Segment III 2.71 × as long as base width. Hypopharynx. Apex of superlingua truncate (Fig. 71F).

Thorax. Dorsal margin of femur with a row of long, stout, pointed setae; one submarginal row of pointed setae as long as those on the dorsal margin (Fig. 72A), ventral margin with short, stout setae and a submarginal row of pointed stout setae, except on hind leg where those on ventral margin are shorter; central area of upper surface with numerous long, stout, blunt setae, shorter on hind femur. Fore tibia with several rows of long simple or feathered setae on ventral margin, dorsal margin with numerous hair-like setae; middle tibia with a row of short, pointed setae and a submarginal row of long, pointed stout setae on ventral margin, outer margin with numerous hair-like setae; hind tibia with a row of short, pointed setae and a submarginal row of long, pointed simple stout setae on ventral margin, lower surface covered with simple, thin setae, outer margin with short and long pointed stout setae and numerous hair-like setae. Tarsal claw hooked with seven subequal denticles, except median much larger (Fig. 72B–C).

Abdomen. Gill I with both lamellae. Gills II–VII with dorsal and ventral lamellae of the same size, or with ventral lamella smaller. Gill I with ~ 15 filaments on each lamella, gills II–VI with ~ 15 filaments on upper lamella, and ~ 10 on lower lamella, gill VII with ~ 15 filaments on each lamella. Filamentous projections on apex, or along entire margin. Posterior margin of terga IX with denticles. Posterior margin of terga VII–VIII with minute denticles. Posterior margin of terga V–VI without denticles. Posterolateral projections present on segments VI–IX.

This species is dedicated to Pauline Sartori, daughter of the first author.

Holotype. • Papua New Guinea, nymph on slide, Gulf Province, Marawaka, Mala, 1400 m, 11.XI.2006, 07°05.664'S, 145°44.467'E, M. Balke and Kinibel col [PNG90], GBIFCH01223165 (ZSM). Paratypes. • Papua New Guinea, one nymph in ethanol (UFVB), one nymph on slide, GBIFCH01223166, 3 nymphs in ethanol, GBIFCH01523451, same data as holotype (MZL). • Papua New Guinea, one nymph on slide, GBIFCH01223167, one nymph in ethanol, GBIFCH01523455, Central Province, Kokoda Trek, 1400 m, I. 2008, 09°14.339'S, 147°40.538'E, Posman col [PNG171] (MZL). • Papua New Guinea, 2 nymphs on slide, GBIFCH01223168, GBIFCH01223169, 3 nymphs in ethanol, GBIFCH01523452, Central Province, Kokoda Trek, 1390 m, I. 2008, 09°00.338'S, 147°44.252'E. Posman col [PNG173] (MZL). • Papua New Guinea, 3 nymphs on slide GBIFCH01223170, GBIFCH01223171, GBIFCH01223172, 4 nymphs in ethanol, GBIFCH01523454, Western Highlands Province, Kundum, 1400 m, 3.III.2007, 05°16.096'S, 144°27.869'E, Kinibel col [PNG142] (MZL). • Papua New Guinea, 3 nymphs on slide, GBIFCH01223173, GBIFCH01223174, GBIFCH01223175, one nymph in ethanol, GBIFCH01523453, Madang Province, Simbai area, 1200 m, 10.III.2007, 05°13.389'S, 144°37.285'E, Kinibel col [PNG152] (MZL). • Papua New Guinea, one nymph on slide, GBIFCH01223176, Madang Province, Simbai area, 1200 m, 11.III.2007, 05°13.333'S, 144°37.611'E, Kinibel col [PNG153] (MZL). • Papua New Guinea, one nymph on slide, GBIFCH01223177, Western Highlands Province, Simbai, 1800–2000 m, 26.II.2007, 05°15.872'S, 144°32.717'E, Kinibel col [PNG134] (MZL).