taxonID	type	description	language	source
49A29F6899385331BB36E82080C2252B.taxon	type_taxon	Type species. Papuanatula (Papuafiliola) stenophylla sp. nov.	en	Kaltenbach, Thomas, Kluge, Nikita J., Gattolliat, Jean-Luc (2025): Redescription of Papuanatula Lugo-Ortiz & McCafferty (Ephemeroptera, Baetidae), with description of a new subgenus and 20 new species. ZooKeys 1227: 159-347, DOI: 10.3897/zookeys.1227.138100
49A29F6899385331BB36E82080C2252B.taxon	etymology	Etymology. The new genus-group name Papuafiliola is formed from the nouns Papua and filiola (most known Latin word for “ little daughter ”). Gender is feminine.	en	Kaltenbach, Thomas, Kluge, Nikita J., Gattolliat, Jean-Luc (2025): Redescription of Papuanatula Lugo-Ortiz & McCafferty (Ephemeroptera, Baetidae), with description of a new subgenus and 20 new species. ZooKeys 1227: 159-347, DOI: 10.3897/zookeys.1227.138100
49A29F6899385331BB36E82080C2252B.taxon	diagnosis	Diagnosis (larval characters). Antenna with each flagellomere symmetric, cylindrical, without brown hypodermal spot (Fig. 133 b) (in contrast to Papuanatula s. str.). Labrum not enlarged and widest at base, dorsally with few submarginal, simple setae (in contrast to feathered setae in Papuanatula s. str.) Mandibles with incisor shortened (Fig. 134 b – e) (in contrast to elongated in Papuanatula s. str.). Labium: Glossae longer than half of paraglossae, with finger-like (distal) portion much longer than triangular (proximal) portion. Labial palp with distomedian projection on 2 nd segment (Fig. 134 h – j) (in contrast to Papuanatula s. str.). Legs: outer side of femur with regular row of long, slender, flattened, parallel-sided setae with blunt apices. Tibia and tarsus with regular row of similar setae. Inner margin of tarsus with distalmost seta not longer or only slightly longer than others. Paraproct without posterior prolongation. Pose of subimaginal gonostyli under larval cuticle in “ Crassolus - type ” (Kluge et al. 2020; Kluge 2021): gonostylus shortened and bent in articulation of 2 nd and 3 rd segments; 2 nd segment directed caudally-medially and not bent; 3 rd segment directed medially-cranially and terminated mediad of 2 nd segment. Besides these characters, both known species of Papuafiliola subgen. nov. have common peculiar characters: body, legs, and marginal ribs of tergalii are covered with minute denticles (Figs 135 a, b, d, 136 k), which form fields of peculiar shape on femora. Eggs of both species with net-like relief (Figs 138 a, b, 145 a – e).	en	Kaltenbach, Thomas, Kluge, Nikita J., Gattolliat, Jean-Luc (2025): Redescription of Papuanatula Lugo-Ortiz & McCafferty (Ephemeroptera, Baetidae), with description of a new subgenus and 20 new species. ZooKeys 1227: 159-347, DOI: 10.3897/zookeys.1227.138100
2D0E5D311DEE532F87CE60ED436663FC.taxon	description	Figs 132, 133, 134, 135, 136, 137	en	Kaltenbach, Thomas, Kluge, Nikita J., Gattolliat, Jean-Luc (2025): Redescription of Papuanatula Lugo-Ortiz & McCafferty (Ephemeroptera, Baetidae), with description of a new subgenus and 20 new species. ZooKeys 1227: 159-347, DOI: 10.3897/zookeys.1227.138100
2D0E5D311DEE532F87CE60ED436663FC.taxon	etymology	Etymology. The species name stenophylla (from στενος — slender, and φυλλον — leaf) refers to theslender tergalii.	en	Kaltenbach, Thomas, Kluge, Nikita J., Gattolliat, Jean-Luc (2025): Redescription of Papuanatula Lugo-Ortiz & McCafferty (Ephemeroptera, Baetidae), with description of a new subgenus and 20 new species. ZooKeys 1227: 159-347, DOI: 10.3897/zookeys.1227.138100
2D0E5D311DEE532F87CE60ED436663FC.taxon	diagnosis	Diagnosis. Larva. The following combination of characters distinguishes P. stenophylla sp. nov. from the other species of Papuanatula (Papuafiliola): femora and abdomen with brown hypodermal maculae; without median protuberances on abdominal terga; tergalii narrow.	en	Kaltenbach, Thomas, Kluge, Nikita J., Gattolliat, Jean-Luc (2025): Redescription of Papuanatula Lugo-Ortiz & McCafferty (Ephemeroptera, Baetidae), with description of a new subgenus and 20 new species. ZooKeys 1227: 159-347, DOI: 10.3897/zookeys.1227.138100
2D0E5D311DEE532F87CE60ED436663FC.taxon	description	Description. Larva (Figs 132 – 136). Cuticular coloration. Head, pronotum and mesonotum mostly pale brownish, with some paler areas. Fore protopteron nearly uniformly pale brownish (Fig. 133 b, e). Metanotum darker brownish medially, colorless laterally. Thoracic pleura pale brownish, sterna mostly colorless. Cuticle of legs mostly colorless, with brownish outer margin and two brownish transverse bands on anterior surface: one close to base and another in apical part; cuticle of these brownish areas serrate (Fig. 135 a, b). Abdominal terga mostly pale brownish; lateral areas of anterior terga paler or colorless; sterna colorless (Fig. 133 c). Cerci uniformly pale brownish. Hypodermal coloration. Anterior side of each femur with wide, contrasting, dark brown macula in middle of area with non-pigmented cuticle (Fig. 136 a – c). Abdomen with sublateral, transverse, contrasting, dark brown maculae between terga (Fig. 132 a, b). Tissues surrounding tracheae of tergalii (main trachea and its branches) with extensive dark brown pigmentation (Fig. 136 d – k). Head. Antenna (Fig. 133 b). As typical for subgenus, with each flagellomere symmetric, cylindrical. Developing turbinate eyes in last instar male larva (Fig. 133 a) with facets equally developed on middle and periphery areas. Labrum (Fig. 134 a) short and widest at base; long setae on dorsal surface simple (not branched), few (2 – 4 on each side) and irregularly situated. Right mandible (Fig. 134 c, e) short, with incisor not elongated; incisor terminating with 4 denticles, most distal denticle shorter than others; other three denticles subequal; mola with seta on proximal corner. Left mandible (Fig. 134 b, d) short, with incisor not elongated; incisor terminating with four denticles, most distal denticle shorter than others; other three denticles subequal; mola with seta on proximal corner. Hypopharynx (Fig. 134 g) with bunch of long, straight, stout, setae-like spines situated on common projection. Maxilla (Fig. 134 f) short and wide; maxillary palp 2 - segmented, as long as galea-lacinia. Labium (Fig. 134 h – j) Paraglossae widest at base, lateral side without concavity; three apical setal rows straight (not bent at apex of paraglossa). Glossa longer than half of paraglossa, with finger-like (distal) portion much longer than triangular (proximal) portion. Ventral side of glossa with several irregularly arranged setae both on triangular and finger-like portions. Labial palp with long, narrow, arched distomedian projection on segment II; segment III with median margin shorter than lateral margin. Thorax. Sterna without protuberances. Terga. Without long setae on midline. Metanotum without hind protoptera or their vestiges. Legs (Figs 135 a – c, 136 a – c). Fore femur widened in proximal part; hind tibia shorter than others. Femur. Outer side of each femur with single regular row of long, slender, flattened, parallel-sided setae with blunt apices. Cuticle of anterior surface of each femur with serrate areas corresponding with brownish pigmentation (see above). Tibia. Patella-tibial suture present on all legs, terminated proximad of inner margin of tibia. Tibia-tarsal condylus turned to anterior side. Anterior side of each tibia with regular row of long, slender, flattened, parallel-sided setae with blunt apices, similar to that on femur, but shorter. Tarsus. Anterior side of each tarsus with regular row of similar, but shorter setae. Posterior side of each tarsus with regular row of few small, stout, pointed setae, most distal of which not longer or slightly longer than others. Claw with row of 9 – 11 subequal denticles and with long, arched seta on posterior side. Abdomen. Terga (Figs 133 c, 135 d). Abdominal terga without long setae on midline. All abdominal terga smooth, without median or submedian elevations or protuberances. Surface of abdominal terga rough, with numerous short sensilla, without long setae; posterior margins of abdominal terga II – X with very small, blunt, dark brown denticles. Posterior margins of abdominal sterna without denticles. Tergalii (Fig. 136 d – k) of abdominal segment I absent; tergalii II – VII subequal, long and narrow. Each tergalius with anal rib longer than costal rib; ribs with very small, irregularly situated denticles on dorsal side. Paraproct (Fig. 133 g) with small, equal denticles on posterior margin, without denticles on median margin, without posterior prolongation. Caudalii (Fig. 133 d). Cerci without swimming setae. Paracercus short, consisting of ~ 15 segments. Pose of subimaginal gonostyli under larval cuticle (Fig. 137 e). In mature larva ready to molt to subimago, subimaginal gonostyli packed under larval cuticle in “ Crassolus - type ” pose (Kluge et al. 2020; Kluge 2021): gonostylus shortened and bent in articulation of 2 nd and 3 rd segments; 2 nd segment directed caudally-medially and not bent; 3 rd segment directed medially-cranially and terminated mediad of 2 nd segment. Subimago. Cuticular coloration (Fig. 137 d). Mesonotum very pale brownish with medioparapsidal sutures colorless, some sutures darker brownish. Thoracic pleura with brown and nearly colorless areas. Sterna nearly colorless. Wing membrane colorless, microtrichia brown. Legs nearly colorless; outer side of femur and base of tibia tinged with pale brownish. Abdomen very pale brownish, cerci colorless. Hypodermal coloration. As in imago. Texture. On all legs of female, terminal tarsomere covered with pointed microlepides only; other tarsomeres covered partly with pointed, partly with blunt microlepides (as in Fig. 143 c, d). Imago. Imago, male (partly molted from subimago). Head ochre. Antennae ochre. Turbinate eyes dull-red, wide, with faceted surfaces round, contiguous. Thorax ochre, equally pale dorsally, laterally, and ventrally. Fore wing with membrane colorless, veins pale ochre, base of RA and costal brace colored with brownish. Pterostigma with 2 – 3 oblique, incomplete cross veins (Fig. 137 a). Legs ochre; femur of each leg with wide brownish band in distal part and with compact, oval, brown macula in proximal 1 / 2 (Fig. 137 c). On middle and hind legs, tarsus with 2 apical spines, on 1 st + 2 nd and 3 rd tarsomeres. Abdomen mostly whitish, terga VII – VIII pale ochre; each tergum I – VIII with pair of small, dark brown sublateral maculae on posterior margin (Fig. 137 b). Cerci pale ochre. Shape of gonostyli unknown. Imago, female. Unknown. Egg (Fig. 138 a, b). Elongate oval. Chorion entirely and evenly covered with ridges forming a net-like relief.	en	Kaltenbach, Thomas, Kluge, Nikita J., Gattolliat, Jean-Luc (2025): Redescription of Papuanatula Lugo-Ortiz & McCafferty (Ephemeroptera, Baetidae), with description of a new subgenus and 20 new species. ZooKeys 1227: 159-347, DOI: 10.3897/zookeys.1227.138100
2D0E5D311DEE532F87CE60ED436663FC.taxon	distribution	Distribution. New Guinea (Fig. 148).	en	Kaltenbach, Thomas, Kluge, Nikita J., Gattolliat, Jean-Luc (2025): Redescription of Papuanatula Lugo-Ortiz & McCafferty (Ephemeroptera, Baetidae), with description of a new subgenus and 20 new species. ZooKeys 1227: 159-347, DOI: 10.3897/zookeys.1227.138100
EA88C84262745D8682259B48D59E7A64.taxon	description	Figs 138, 139, 140, 141, 142, 143, 144, 145	en	Kaltenbach, Thomas, Kluge, Nikita J., Gattolliat, Jean-Luc (2025): Redescription of Papuanatula Lugo-Ortiz & McCafferty (Ephemeroptera, Baetidae), with description of a new subgenus and 20 new species. ZooKeys 1227: 159-347, DOI: 10.3897/zookeys.1227.138100
EA88C84262745D8682259B48D59E7A64.taxon	etymology	Etymology. The species name tuberculata refers to the unpaired tubercles on abdominal terga of the larva (Fig. 138 c).	en	Kaltenbach, Thomas, Kluge, Nikita J., Gattolliat, Jean-Luc (2025): Redescription of Papuanatula Lugo-Ortiz & McCafferty (Ephemeroptera, Baetidae), with description of a new subgenus and 20 new species. ZooKeys 1227: 159-347, DOI: 10.3897/zookeys.1227.138100
EA88C84262745D8682259B48D59E7A64.taxon	diagnosis	Diagnosis. Larva. The following combination of characters distinguishes P. tuberculata sp. nov. from the other species of Papuafiliola subgen. nov.: absence of brown hypodermal maculae on femora and abdomen; presence of median protuberances on abdominal terga; tergalii rather wide.	en	Kaltenbach, Thomas, Kluge, Nikita J., Gattolliat, Jean-Luc (2025): Redescription of Papuanatula Lugo-Ortiz & McCafferty (Ephemeroptera, Baetidae), with description of a new subgenus and 20 new species. ZooKeys 1227: 159-347, DOI: 10.3897/zookeys.1227.138100
EA88C84262745D8682259B48D59E7A64.taxon	description	Description. Larva (Figs 138 – 142). Cuticular coloration. Head, pronotum and mesonotum mostly pale brownish, with some paler areas. Fore protopteron pale brownish, with narrow paler lines along concave veins (Fig. 139 a, c). Metanotum darker brownish medially, colorless laterally. Thoracic pleura pale brownish, sterna colorless. Cuticle of legs mostly colorless, with brownish outer margin and two brownish transverse bands on anterior surface: one close to base and another in apical part; cuticle of these brownish areas serrate (Figs 139 b, 141 a, b). Abdominal terga mostly pale brownish, with lateral areas paler or colorless; tergum VI more or less paler than others; sterna colorless (Figs 138, 139 d). Cerci uniformly pale brownish. Hypodermal coloration. Not expressed (at least in penultimate larval instar). Tissues surrounding tracheae of tergalii (main trachea and its branches) with brown pigmentation (Fig. 142 d, e). Head. Antenna (Fig. 142 c) with each flagellomere symmetric, cylindrical. Developing turbinate eyes in last instar male larva. Unknown. Labrum (Fig. 140 a) short and widest at base; long setae on dorsal surface simple (not branched), constitute two pairs: submedian pair and one more pair laterad of them. Right mandible (Fig. 140 c) short, with incisor not elongated; incisor terminating with four denticles, among which most distal denticle shorter than others; other three denticles subequal; mola with seta on proximal corner. Left mandible (Fig. 140 b) short, with incisor not elongated; incisor terminating with four denticles, among which most distal denticle shorter than others; other three denticles subequal; mola with seta on proximal corner. Hypopharynx (Fig. 140 f, g) with bunch of long, straight, stout, setae-like spines situated on common projection. Maxilla (Fig. 140 d, e) short and wide; maxillary palp 2 - segmented, as long as galea-lacinia. Labium (Fig. 140 h – j). Paraglossae widest at base, lateral side with slight concavity near base; three apical setal rows parallel to apical-lateral margin only (not bent parallel to median margin). Glossa longer than half of paraglossa, with finger-like (distal) portion much longer than triangular (proximal) portion. Ventral side of glossa with several irregularly arranged setae both on triangular and finger-like portions. Labial palp with long, narrow, arched distomedian projection on segment II; segment III with median margin shorter than lateral margin. Thorax. Sterna without protuberances. Terga. Without long setae on midline; without protuberances. Metanotum without hind protoptera or their vestiges. Legs (Figs 139 b, 141 a, b). Fore femur widened in proximal part; hind tibia shorter than others. Femur. Outer side of each femur with single regular row of long, slender, flattened, parallel-sided setae with blunt apices. Cuticle of anterior surface of each femur with serrate areas corresponding with brownish pigmentation (see above). Tibia. Patella-tibial suture present on all legs, terminated proximad of inner margin of tibia. Tibia-tarsal condylus turned to anterior side. Anterior side of each tibia with regular row of long, slender, flattened, parallel-sided setae with blunt apices, similar to that on femur, but shorter. Tarsus. Anterior side of each tarsus with regular row of similar, but shorter setae. Posterior side of each tarsus with regular row of few small, stout, pointed setae, most distal of which not longer or slightly longer than others. Claw with row of 8 – 10 subequal denticles and with long, arched seta on posterior side. Abdomen. Terga (Figs 138, 139 d, 142 a) without long setae on midline (e. g., in contrast to P. obscura sp. nov.). Each abdominal tergum I – IX with slightly expressed median ridge elevating toward posterior margin. Surface of abdominal terga serrate, with numerous short sensilla; posterior margins of abdominal terga II – X with very small, blunt, dark brown denticles. Abdominal sterna without serrations and denticles. Tergalii (Fig. 142 d, e) of abdominal segment I absent; tergalii II – VII subequal, oval. Each tergalius with anal rib longer than costal rib; ribs with very small, irregularly situated denticles on dorsal side. Paraproct (Fig. 142 b) with many small, equal denticles on median and posterior margins, without posterior prolongation. Caudalii (Fig. 139 d). Cerci without swimming setae. Paracercus short, consisting of ~ 12 segments. Pose of subimaginal gonostyli under larval cuticle. Unknown. Subimago. Cuticular coloration. Pronotum partly brownish. Mesonotum pale brown with medioparapsidal suture colorless, other sutures darker brown (Fig. 143 a). Meso- and metathoracic pleura and sterna with colorless, pale brownish and dark brown areas (Fig. 143 b). Cuticle of wings colorless, with microtrichiae brownish. Legs nearly colorless, with pale brown bordering on femur and base of tibia (as in Fig. 36 d). Abdomen very pale brownish. Cerci colorless with setae brownish. Hypodermal coloration. As in imago. Texture. On all legs of female, terminal tarsomere covered with pointed microlepides only; other tarsomeres covered partly with pointed, partly with blunt microlepides (Fig. 143 c, d). Imago. Imago, male. Unknown. Imago, female (Fig. 144 a – f). Head ochre with brown. Antennae ochre, distal parts of scapus and pedicellum brown. Thorax brown with ochre, equally dark dorsally, laterally, and ventrally. Fore wing with membrane mostly colorless, veins pale ochre; base of wing, including costal brace contrastingly pigmented with brown and ochre. Pterostigma with three or four oblique, incomplete cross veins (Fig. 144 f). Legs ochre; femur of each leg with wide reddish-brown band in distal part; femur of fore leg, besides this, with longitudinal reddish-brown macula occupying most part of proximal 1 / 2 (Fig. 144 c – e). On each leg, tarsus with two apical spines: on 2 nd and 3 rd tarsomeres on fore leg, on 1 st + 2 nd and 3 rd tarsomeres on middle and hind legs. Each abdominal tergum I – IX brown with paler ochre areas; tergum X brownish ochre; sterna ochre (Fig. 144 a, b). Cerci pale ochre. Egg (Fig. 145 a – e). Elongate oval. Chorion entirely and evenly covered with ridges forming a net-like relief.	en	Kaltenbach, Thomas, Kluge, Nikita J., Gattolliat, Jean-Luc (2025): Redescription of Papuanatula Lugo-Ortiz & McCafferty (Ephemeroptera, Baetidae), with description of a new subgenus and 20 new species. ZooKeys 1227: 159-347, DOI: 10.3897/zookeys.1227.138100
EA88C84262745D8682259B48D59E7A64.taxon	distribution	Distribution. New Guinea (Fig. 148).	en	Kaltenbach, Thomas, Kluge, Nikita J., Gattolliat, Jean-Luc (2025): Redescription of Papuanatula Lugo-Ortiz & McCafferty (Ephemeroptera, Baetidae), with description of a new subgenus and 20 new species. ZooKeys 1227: 159-347, DOI: 10.3897/zookeys.1227.138100
A8CB813B5D5B586EBAB13E874CF478B4.taxon	description	Figs 26, 27, 28, 29, 30, 31	en	Kaltenbach, Thomas, Kluge, Nikita J., Gattolliat, Jean-Luc (2025): Redescription of Papuanatula Lugo-Ortiz & McCafferty (Ephemeroptera, Baetidae), with description of a new subgenus and 20 new species. ZooKeys 1227: 159-347, DOI: 10.3897/zookeys.1227.138100
A8CB813B5D5B586EBAB13E874CF478B4.taxon	etymology	Etymology. The species is dedicated to Michael Balke (ZSM / SNSB), the collector of an important part of the material used in this study.	en	Kaltenbach, Thomas, Kluge, Nikita J., Gattolliat, Jean-Luc (2025): Redescription of Papuanatula Lugo-Ortiz & McCafferty (Ephemeroptera, Baetidae), with description of a new subgenus and 20 new species. ZooKeys 1227: 159-347, DOI: 10.3897/zookeys.1227.138100
A8CB813B5D5B586EBAB13E874CF478B4.taxon	diagnosis	Diagnosis. Larva. The following combination of characters distinguishes P. balkei sp. nov. from other species of Papuanatula s. str.: body dorsally with irregular row of long, fine, simple setae along midline; abdominal terga without protuberances; femur proximally with wedge-shaped blank, overlaid with scattered brown color; abdominal terga brown, laterally darker, terga V, VI, and X brighter; tergalii with pigmented tracheation; paracercus with seven or eight segments; abdominal terga with triangular, apically rounded denticles on posterior margin; small scattered scales on abdominal terga elongate, slightly trapezoid.	en	Kaltenbach, Thomas, Kluge, Nikita J., Gattolliat, Jean-Luc (2025): Redescription of Papuanatula Lugo-Ortiz & McCafferty (Ephemeroptera, Baetidae), with description of a new subgenus and 20 new species. ZooKeys 1227: 159-347, DOI: 10.3897/zookeys.1227.138100
A8CB813B5D5B586EBAB13E874CF478B4.taxon	description	Description. Larva (Figs 26 – 31). Body length 4.2 – 5.6 mm, cerci much longer than body length (~ 1.5 ×). Cuticular coloration (Fig. 26 a – c). Head, thorax and abdomen dorsally brown; thorax with complex pattern; abdominal segments laterally darker, V, VI, and X brighter. Femur proximally with wedge-shaped blank, overlaid with scattered brown color; medial area grey-brown, distal area yellow-brown to grey-brown; tibia grey; tarsus grey, distally brown. Head, thorax and abdominal segment I ventrally ecru, protuberances of thoracic sterna brown; abdominal segments II – X ventrally pale brown, darker toward end of abdomen. Cerci grey-brown. Hypodermal coloration. Each abdominal tergum I – IX with wide dark brown transverse band close to anterior margin and with narrower dark brown transverse band close to posterior margin (Fig. 26 a). Head (Figs 26 b, 29 h). Dorsally with irregular row of long, fine, simple setae along midline. Antenna. Length ~ 1.5 × head length. Otherwise, as typical for subgenus. Developing turbinate eyes in last instar male larva (Fig. 29 h) ovoid, with large distance to each other. Labrum (Fig. 27 a, b). Length 0.5 × maximum width, laterally convex. Dorsal, sub-marginal arc with 24 – 29 feathered setae. Right mandible (Fig. 27 d, e). Margin between prostheca and mola with row of minute denticles. Otherwise, as typical for subgenus. Left mandible (Fig. 27 f, g). Margin between prostheca and mola with row of minute denticles. Otherwise, as typical for subgenus. Hypopharynx (Fig. 27 c). As typical genus. Maxilla (Fig. 28 c, d). Maxillary palp subequal in length to galea-lacinia; palp segment II slightly longer than segment I. Otherwise, as typical for genus. Labium (Fig. 28 a, b). As typical for genus. Paraglossa dorsally with two spine-like setae near inner, distolateral margin. Labial palp with segment I subequal in length to segments II and III combined. Segment II with minute distomedial protuberance, dorsally with row of four spine-like setae near outer, distolateral margin. Segment III slightly pentagonal, pointed; 0.7 × length of segment II; inner dorsal margin with few feathered setae. Thorax. Sterna. With small protuberances on sides of prosternum and close to openings of mesothoracic and metathoracic sternal apodemes (as in Fig. 108 a). Terga (Fig. 26 b) without protuberances; with irregular row of long, fine, simple setae along midline. Metanotum without hind protoptera or their vestiges. Legs (Fig. 29 a – g). Ratio of leg segments: fore leg 0.8: 1.0: 0.4: 0.1, middle leg 0.9: 1.0: 0.3: 0.1 and hind leg 1.0: 1.0: 0.4: 0.2. Femur. Length ~ 3.7 × maximum width. Claw with one row of 5 – 7 denticles and one or sometimes two posterior setae. Abdomen. Terga (Figs 30 a, 31 a – c) with irregular row of long, fine, simple setae along midline. Terga without protuberances, terga I – IV with slight, paired medioposterior elevations. Posterior margin of terga: I with rudimentary denticles, II – IX with triangular, apically rounded denticles. Surface with scattered small, elongate, slightly trapezoid, striated scales. Tergalii (Fig. 30 d, e). Broad ovoid, tracheation well pigmented; margins smooth, with few short, fine, simple setae. Paraproct (Fig. 30 f). Posterior margin with prolongation and row of minute denticles. Caudalii (Fig. 30 b, c). Cerci apart from basal and distal part with 1 – 7 swimming setae per segment, initially increasing and then again decreasing toward distal part. Paracercus with seven or eight segments. Pose of subimaginal gonostyli under larval cuticle. Unknown. Subimago. Unknown. Imago. Unknown. Egg. Unknown.	en	Kaltenbach, Thomas, Kluge, Nikita J., Gattolliat, Jean-Luc (2025): Redescription of Papuanatula Lugo-Ortiz & McCafferty (Ephemeroptera, Baetidae), with description of a new subgenus and 20 new species. ZooKeys 1227: 159-347, DOI: 10.3897/zookeys.1227.138100
A8CB813B5D5B586EBAB13E874CF478B4.taxon	distribution	Distribution. New Guinea (Fig. 146).	en	Kaltenbach, Thomas, Kluge, Nikita J., Gattolliat, Jean-Luc (2025): Redescription of Papuanatula Lugo-Ortiz & McCafferty (Ephemeroptera, Baetidae), with description of a new subgenus and 20 new species. ZooKeys 1227: 159-347, DOI: 10.3897/zookeys.1227.138100
3EF770306D6A5AF7B1A59EAC62EA7461.taxon	description	Figs 1, 2, 3, 4, 5, 6	en	Kaltenbach, Thomas, Kluge, Nikita J., Gattolliat, Jean-Luc (2025): Redescription of Papuanatula Lugo-Ortiz & McCafferty (Ephemeroptera, Baetidae), with description of a new subgenus and 20 new species. ZooKeys 1227: 159-347, DOI: 10.3897/zookeys.1227.138100
3EF770306D6A5AF7B1A59EAC62EA7461.taxon	diagnosis	Diagnosis. Larva. The following combination of characters distinguishes P. bessa from other species of Papuanatula s. str.: body dorsally with irregular row of long, fine, simple setae along midline; metanotum and abdominal terga I – V with medioposterior, broad, paired humps; abdominal terga II – IV dark brown, with brighter oblong markings; tergum V yellow brown with T-shaped dark brown marking (Fig. 1 a; Lugo-Ortiz and McCafferty 1999: fig. 5); femur brownish, with wedge-shaped blank in proximal 1 / 2; paracercus with eight segments.	en	Kaltenbach, Thomas, Kluge, Nikita J., Gattolliat, Jean-Luc (2025): Redescription of Papuanatula Lugo-Ortiz & McCafferty (Ephemeroptera, Baetidae), with description of a new subgenus and 20 new species. ZooKeys 1227: 159-347, DOI: 10.3897/zookeys.1227.138100
3EF770306D6A5AF7B1A59EAC62EA7461.taxon	description	Description. Larva (Figs 1 a – 5 h). Body length 3.8 – 4.9 mm, cerci much longer than body length (~ 1.5 ×). Cuticular coloration. Head, pronotum, mesonotum, and metanotum brownish, with darker and paler areas; fore protoptera with narrow paler lines corresponding to all longitudinal and intercalary veins (Fig. 2 c). Thoracic pleura brownish, sterna with brownish and colorless areas. Cuticle of femur with wedge-shape colorless blank on proximal 1 / 2 and less distinct blank occupying most part of distal 1 / 2; other surface of femur brownish, apex bordered with darker brown (Figs 2 d – f, 4 a). Tibia and tarsus brownish (Fig. 2 d – f). Abdominal terga mostly brownish, with lateral areas paler; terga V – VI more or less paler than others; sterna mostly colorless (Figs 1 a, 2 g). Cerci uniformly pale brownish. Hypodermal coloration. Legs without hypodermal markings. Abdominal terga with extensive, contrasting, dark brown markings of characteristic shape (Figs 1 a, 2 a). Tissues surrounding tracheae of tergalii partly with brown pigmentation. Head. Dorsally with row of long, fine, simple setae along midline (as in Figs 26 b, 65 b). Antenna (Fig. 2 a, b). Length ~ 1.5 × head length; otherwise, typical for subgenus. Developing turbinate eyes in last instar male larva. With larger facets in middle and smaller facets on periphery (as in Fig. 66 h). Labrum (Fig. 3 a, b) Length 0.5 × maximum width, laterally convex; widened distally; 30 – 35 long, feathered setae on dorsal surface forming integral, regular transverse row; each seta consists of stout stem and numerous long processes on both sides. Right mandible (Fig. 3 e). Incisor sharply pointed, with two denticles near base. Kinetodontium terminated with three denticles, with distal denticle longest. Otherwise, as typical for subgenus. Left mandible (Fig. 3 f). Incisor sharply pointed, with two denticles near base. Kinetodontium terminated with three denticles, with distal denticle longest. Otherwise, as typical for subgenus. Hypopharynx (Fig. 3 d). As typical for genus. Maxilla (Fig. 3 g). Maxillary palp as long as galea-lacinia. Otherwise, as typical for genus. Labium (Fig. 3 c). Paraglossae widened near middle, with lateral side forming concavity in proximal part; three apical setal rows sharply bent. Glossa as long as half of paraglossa, with finger-like (distal) portion as long as triangular (proximal) portion. Glossa with several long setae at apex and one long seta near middle of ventral side. Labial palp without distomedian projection on segment II; segment III with median margin longer than lateral margin. Thorax. Sterna. With small protuberances on sides of prosternum and close to openings of mesothoracic and metathoracic sternal apodemes (as in Fig. 108 a). Terga with irregular row of long, fine, soft, colorless, simple setae along midline (as in Figs 26 b, 65 b). Metanotum with medioposterior broad, paired elevations. Metanotum without hind protoptera or their vestiges. Legs (Figs 2 d – f, 4 a – e). Fore femur slightly widened in proximal part; hind tibia shorter than others. Ratio of leg segments: fore leg 0.8: 1.0: 0.3: 0.2, middle leg 0.9: 1.0: 0.3: 0.2 and hind leg 1.0: 1.0: 0.3: 0.2. Femur. Length ~ 4 × maximum width. Outer side of each femur with single regular row of long, hair-like setae bearing numerous fine, short branches on all sides (as in Figs 41 g, 68 b). Tibia. Patella-tibial suture present on all legs, terminated near middle of inner margin of tibia. Tibia-tarsal condylus (originally located on outer side) turned to anterior side. Anterior (originally outer-anterior) side of each tibia with regular row of setae similar to that on femur. Tarsus. Anterior (originally outer) side of each tarsus with regular row of similar, but smaller (shorter and narrower) setae. Posterior (originally inner) side of each tarsus with regular row of short, stout, oval setae (looking pointed in profile) and one much longer, thinner, pointed seta distad of them. Claw with row of six or seven short denticles and one somewhat larger denticle distad of them; long, arched posterior seta on posterior side near distal denticle. Abdomen. Terga (Fig. 5 a – g). Long, fine, soft, colorless setae irregularly situated along midline of all abdominal terga (as in Figs 26 b, 65 b). Each abdominal tergum with pair of submedian protuberances, more prominent on terga I – IV and less prominent to absent on more posterior terga; each protuberance with compact group of short, blunt, parallel-sided, colorless scales with round sockets. Other surface with fewer, sparse scales of such kind. Posterior margins of abdominal terga partly without denticles, partly with short, blunt denticles separated by spaces. Tergalii (Figs 2 g, 4 f, g) of abdominal segment I absent; tergalii II – VII subequal, oval. Each tergalius with costal and anal ribs narrow, present on proximal part of tergalius only. Tracheation poorly developed or absent; margins smooth, with short, fine, simple setae. Paraproct (Fig. 5 h) with posterior prolongation bent toward bases of caudalii, with many small, equal denticles on median-posterior margin. Caudalii (Fig. 4 h, i) without swimming setae; vestiges of swimming setae present on distal part of cerci. Paracercus short, consisting of ~ 7 – 10 segments. Pose of subimaginal gonostyli under larval cuticle. In mature larva ready to molt to subimago, subimaginal gonostyli packed under larval cuticle in ’ Labiobaetis - type’ pose: 2 nd segments directed medially and bent proximally; 3 rd segment directed medially (as continuation of 2 nd segment) and narrowed apically, being deformed corresponding to space between subimaginal styliger and larval cuticle (Fig. 6 d). Subimago. Texture. On all legs of both sexes, each tarsomere covered mostly with blunt microlepides, with pointed microlepides near apex (as in Fig. 70 i). Imago. Imago, male. Unknown. Following features can be revealed based on subimaginal structure: Turbinate eyes narrow and cylindrical, with stem reddish and faceted surface small, round, black. Legs ochre, without brown hypodermal markings. Abdomen with characteristic hypodermal coloration forming brown and ochre areas (Fig. 6 a – c). Imago, female. Unknown. Judging by description of subimago (Lugo-Ortiz and McCafferty 1999: 64), hypodermal coloration of abdominal terga as in male. Egg. Unknown.	en	Kaltenbach, Thomas, Kluge, Nikita J., Gattolliat, Jean-Luc (2025): Redescription of Papuanatula Lugo-Ortiz & McCafferty (Ephemeroptera, Baetidae), with description of a new subgenus and 20 new species. ZooKeys 1227: 159-347, DOI: 10.3897/zookeys.1227.138100
3EF770306D6A5AF7B1A59EAC62EA7461.taxon	distribution	Distribution. New Guinea (Fig. 146).	en	Kaltenbach, Thomas, Kluge, Nikita J., Gattolliat, Jean-Luc (2025): Redescription of Papuanatula Lugo-Ortiz & McCafferty (Ephemeroptera, Baetidae), with description of a new subgenus and 20 new species. ZooKeys 1227: 159-347, DOI: 10.3897/zookeys.1227.138100
A7FE8EA98D5758D89C9B33EA0E6025CB.taxon	description	Figs 7, 8, 9, 10, 11, 12, 13	en	Kaltenbach, Thomas, Kluge, Nikita J., Gattolliat, Jean-Luc (2025): Redescription of Papuanatula Lugo-Ortiz & McCafferty (Ephemeroptera, Baetidae), with description of a new subgenus and 20 new species. ZooKeys 1227: 159-347, DOI: 10.3897/zookeys.1227.138100
A7FE8EA98D5758D89C9B33EA0E6025CB.taxon	diagnosis	Diagnosis. Larva. The following combination of characters distinguishes P. copis from other species of Papuanatula s. str.: body dorsally without row of long, fine, simple setae along midline; metanotum and abdominal terga I – VIII medially with conspicuous, long protuberances, with long, fine points, oriented dorsoposteriad, on abdominal segment (s) IX (X) vestigial; labial palp segment II without distomedial protuberance; segment III globular; femur with angulate blank in basal part; paracercus vestigial.	en	Kaltenbach, Thomas, Kluge, Nikita J., Gattolliat, Jean-Luc (2025): Redescription of Papuanatula Lugo-Ortiz & McCafferty (Ephemeroptera, Baetidae), with description of a new subgenus and 20 new species. ZooKeys 1227: 159-347, DOI: 10.3897/zookeys.1227.138100
A7FE8EA98D5758D89C9B33EA0E6025CB.taxon	description	Description. Larva (Figs 7 – 13). Body length 6.0 – 6.5 mm, cerci much longer than body length (~ 2 ×). Cuticular coloration (Figs 7 a, b, 8 a – c). Head, thorax and abdomen dorsally brown to dark brown. Thorax with complex markings. Abdominal segments I – VI (– IX) anteriorly and laterally darker, VII – IX at least laterally darker. Head and thorax ventrally ecru; abdomen ventrally pale yellow-brown, laterally with brown markings. Legs yellow-brown to brown, femur with angulate blank in basal part. Caudalii yellow-brown. Hypodermal coloration. Abdominal segments I – IX dorsally with transverse band on posterior margins (Fig. 8 a). Head. Antenna (Fig. 10 e). Length ~ 1.5 × head length. As typical for subgenus. Developing turbinate eyes in last instar male larva (Fig. 10 e) rather narrow, with big distance from each other. Labrum (Fig. 9 a, b). Length 0.4 × maximum width, laterally slightly angulate. Dorsal, sub-marginal arc with> 50 densely articulated, feathered setae (30 – 35 setae according to Lugo-Ortiz and McCafferty 1999: 65). Right mandible (Fig. 9 d, e). Margin between prostheca and mola smooth, without denticles. Otherwise, as typical for subgenus. Left mandible (Fig. 9 f, g). Margin between prostheca and mola, smooth, without denticles. Subtriangular process often undeveloped. Otherwise, as typical for subgenus. Hypopharynx (Fig. 9 c). As typical for genus. Maxilla (Fig. 10 c, d). Maxillary palp slightly longer than galea-lacinia; palp segment II slender, partly sclerotized, approx. as long as segment I, segment I thicker. Otherwise, as typical for genus. Labium (Fig. 10 a, b). As typical for genus. Paraglossa with two spine-like setae near inner margin. Labial palp with segment I 0.8 × length of segments II and III combined. Segment II without distomedial protuberance, dorsally with row of 2 – 5 spine-like setae near outer, distolateral margin. Segment III globular, slightly pointed, 0.7 × length of segment II; inner dorsal margin with few feathered setae. Thorax. Sterna. Protuberances poorly developed. Terga (Figs 7 a, b, 8 a, b, 13 b). Metanotum medially with conspicuous, long protuberance, oriented dorsoposteriad. Immature larva with short, acute, posteromedial protuberance on pro- and mesonotum. Legs (Fig. 11 a – h). Ratio of leg segments: fore leg 0.9: 1.0: 0.3: 0.1, middle leg 0.9: 1.0: 0.3: 0.1 and hind leg 1.1: 1.0: 0.3: 0.1. Femur. Length ~ 3 × maximum width. Claw with one row of seven or eight denticles and one posterior seta. Abdomen. Terga (Figs 12 a, b, 13 a). Abdominal terga I – VIII medially with conspicuous, long protuberances, with long, fine points, oriented dorsoposteriad, on abdominal segment (s) IX (X) vestigial. Posterior margin of terga I – IX with small, triangular, pointed denticles. Surface with scattered small, conical, apically rounded scales. Tergalii (Fig. 11 i, j) ovoid, tracheation developed; with brown pigmentation in middle area; margins with minute serration and many short, fine, simple setae. Paraproct (Fig. 12 e – g). Posterior margin with prolongation and with row of many minute denticles. Caudalii (Fig. 12 c, d). Cerci without swimming setae; sometimes few vestigial swimming setae or insertions still present. Paracercus vestigial. Pose of subimaginal gonostyli under cuticle. Unknown. Subimago. Unknown. Imago. Unknown. Egg. Unknown.	en	Kaltenbach, Thomas, Kluge, Nikita J., Gattolliat, Jean-Luc (2025): Redescription of Papuanatula Lugo-Ortiz & McCafferty (Ephemeroptera, Baetidae), with description of a new subgenus and 20 new species. ZooKeys 1227: 159-347, DOI: 10.3897/zookeys.1227.138100
A7FE8EA98D5758D89C9B33EA0E6025CB.taxon	distribution	Distribution. New Guinea (Fig. 146).	en	Kaltenbach, Thomas, Kluge, Nikita J., Gattolliat, Jean-Luc (2025): Redescription of Papuanatula Lugo-Ortiz & McCafferty (Ephemeroptera, Baetidae), with description of a new subgenus and 20 new species. ZooKeys 1227: 159-347, DOI: 10.3897/zookeys.1227.138100
835CD3DA6248573393280208AFAD25D1.taxon	description	Figs 32, 33, 34, 35, 36, 37	en	Kaltenbach, Thomas, Kluge, Nikita J., Gattolliat, Jean-Luc (2025): Redescription of Papuanatula Lugo-Ortiz & McCafferty (Ephemeroptera, Baetidae), with description of a new subgenus and 20 new species. ZooKeys 1227: 159-347, DOI: 10.3897/zookeys.1227.138100
835CD3DA6248573393280208AFAD25D1.taxon	etymology	Etymology. The species name cyclopomontana refers to Cyclops Mountain, at which foot this species was collected.	en	Kaltenbach, Thomas, Kluge, Nikita J., Gattolliat, Jean-Luc (2025): Redescription of Papuanatula Lugo-Ortiz & McCafferty (Ephemeroptera, Baetidae), with description of a new subgenus and 20 new species. ZooKeys 1227: 159-347, DOI: 10.3897/zookeys.1227.138100
835CD3DA6248573393280208AFAD25D1.taxon	diagnosis	Diagnosis. Larva. The following combination of characters distinguishes P. cyclopomontana sp. nov. from other species of Papuanatula s. str.: body dorsally with irregular row of long, fine, simple setae along midline; abdominal terga without protuberances; paraglossa with three straight setal rows (not bent at apex of paraglossa); femur basally with wedge-shaped blank and less contrasting blank on distal ¹ ⁄ 3; tergalii with extensive, brown pigmentation; paracercus with 6 – 8 segments; abdominal terga II – IX with various denticles, from long and pointed to short and blunt.	en	Kaltenbach, Thomas, Kluge, Nikita J., Gattolliat, Jean-Luc (2025): Redescription of Papuanatula Lugo-Ortiz & McCafferty (Ephemeroptera, Baetidae), with description of a new subgenus and 20 new species. ZooKeys 1227: 159-347, DOI: 10.3897/zookeys.1227.138100
835CD3DA6248573393280208AFAD25D1.taxon	description	Description. Larva (Figs 33 – 35). Cuticular coloration. Head, pronotum, mesonotum and metanotum brownish, with darker and paler areas; fore protoptera nearly uniformly brown (Fig. 32 c, d). Thoracic pleura brownish, sterna mostly colorless. Cuticle of femur mostly brownish, with clearly outlined wedge-shape blank on proximal ¹ ⁄ 3 and less contrasting blank on distal ¹ ⁄ 3; apex of femur bordered with darker brown (Fig. 32 e – g). Tibia and tarsus mostly brownish (Fig. 32 e – g). Abdominal terga either mostly brownish, or with brown anterior margin and paler remainder part; terga V and VI paler than others (Fig. 32 a). Sterna mostly colorless. Cerci uniformly pale brownish. Hypodermal coloration. Judging by hypodermal coloration of male imago and female subimagines, legs without hypodermal markings; each abdominal tergum I – IX with dark brown transverse band close to posterior margin (Fig. 36 b, c, g). Tissues surrounding tracheae of tergalii (main trachea and its branches) with extensive brown pigmentation (Fig. 32 a). Head. Dorsally with irregular row of long, fine, simple setae along midline. Antenna (Fig. 32 d). Length ~ 1.5 × head length. As typical for subgenus. Developing turbinate eyes in last instar male larva (Fig. 32 d) with facets equally developed on middle and peripheral areas. Labrum (Fig. 33 a) widened distally; long, feathered setae on dorsal surface numerous and forming integral, regular transverse row. Right mandible. As typical for subgenus. Left mandible. As typical for subgenus. Hypopharynx (Fig. 33 b). As typical for genus. Maxilla (Fig. 33 c). Maxillary palp shorter than galea-lacinia. Otherwise, as typical for genus. Labium (Fig. 33 d). Paraglossae with proximal 1 / 2 nearly parallel-sided; three apical setal rows straight (not bent at apex of paraglossa). Glossa as long as half of paraglossa, with finger-like (distal) portion as long as triangular (proximal) portion. Labial palp without distomedial projection on segment II; segment III with median margin longer than lateral margin. Thorax. Sterna. With small protuberances on sides of prosternum and close to openings of mesothoracic and metathoracic sternal apodemes (as Fig. 108 a). Terga without protuberances. Long, fine, soft, colorless setae irregularly situated along midline of all terga (as in Fig. 65 b). Metanotum without hind protoptera or their vestiges. Legs (Figs 32 e – g, 34 c – e). Fore femur widened in proximal part; hind tibia shorter than others. Femur. Outer side of each femur with single regular row of long, hair-like setae bearing numerous fine, short branches on all sides (as in Figs 41 g, 68 b). Tibia. Patella-tibial suture present on all legs, terminated near middle of inner margin of tibia. Tibia-tarsal condylus turned to anterior side. Anterior side of each tibia with regular row of setae similar to that on femur. Tarsus. Anterior side of each tarsus with regular row of similar, but smaller (shorter and narrower) setae. Posterior side of each tarsus with regular row of short, stout, oval setae (looking pointed in profile) and one much longer, thinner, pointed seta distad of them. Claw with row of 5 – 8 short denticles and one somewhat larger denticle distad of them; long, arched, posterior seta. Abdomen. Terga (Figs 32 a, 35 a, b). Long, fine, soft, colorless setae irregularly situated along midline of all abdominal terga (as in Fig. 65 b). Abdominal terga without dorsal unpaired or paired protuberances, only with slightly expressed unpaired, median elevations. Abdominal terga with small, roundish scales with small sockets and radial striation (visible in dry condition, but not in Canada balsam). Posterior margins of abdominal terga II – IX with various denticles, from long and pointed to short and blunt, more numerous and long on middle terga, few and short on terga II and IX. Posterior margin of tergum X with smaller, blunt denticles. Tergalii (Fig. 32 a, h) II – VII subequal, oval. Each tergalius with costal and anal ribs narrow, smooth, present on proximal 1 / 2 of tergalius only. Paraproct (Fig. 35 c) with regular row of small, pointed, equal denticles on median margin, without posterior prolongation. Caudalii (Fig. 34 a, b) without swimming setae; vestiges of swimming setae present on distal part of cerci. Paracercus short, consisting of ~ 6 – 8 segments. Pose of subimaginal gonostyli under larval cuticle. Unknown. Subimago. Cuticular coloration. Pronotum and prosternum partly brown (as in Fig. 60 f). Mesonotum pale brown with medioparapsidal suture colorless, other sutures darker brown (Fig. 36 f). Meso- and metathoracic pleura and sterna with colorless, pale brownish and dark brown areas (Fig. 36 e). Cuticle of wings colorless, with microtrichiae brownish. Legs nearly colorless, with pale brown bordering on femur and base of tibia (Fig. 36 d). Abdomen diffusely colored with very pale brownish in distal part. Hypodermal coloration. As in imago. Texture. On all legs of both sexes, each tarsomere covered mostly with blunt microlepides, with pointed microlepides near apex (as in Fig. 70 i). Imago. Imago, male. Head pale ochre. Antennae ochre. Turbinate eyes yellow, widened apically. Thorax ochre, equally pale dorsally, laterally, and ventrally. Fore wing with membrane colorless, veins ochre. Pterostigma with three or four incomplete, oblique cross veins. Legs ochre (Fig. 36 b, c). Abdominal terga mostly ochre, terga II – IV with median part pale brown; each tergum I – IX with darker brown transverse band close to posterior margin; sterna ochre (Fig. 36 g). Genitalia (Fig. 37). Sterno-styligeral muscle absent. Each unistyliger nearly equally wide at base and at apex, with median margin concave. At lateral side of gonostylus, its 1 st segment roundly-convex and separated from 2 nd segment by concavity; at median side of gonostylus, 1 st segment gradually turns to 2 nd segment. 2 nd segment equally wide all over its length. Third (terminal) segment of gonostylus nearly as wide as 2 nd, with length slightly exceeding width. Penial bridge without projection between unistyligers. Gonovectes dark brown. Each gonovectis parabolic, with lateral (basal) and median (apical) portions equally long, apex bent medially. Imago, female. Head and thorax dorsally ochre-brownish. Abdomen mostly ochre, terga and sterna with brownish markings, each tergum I – IX with dark brown transverse band close to posterior margin (Fig. 36 h, i). Coloration of legs, wings, and cerci as in male. Egg (Fig. 37 b). Irregularly oval. Chorion without regular relief.	en	Kaltenbach, Thomas, Kluge, Nikita J., Gattolliat, Jean-Luc (2025): Redescription of Papuanatula Lugo-Ortiz & McCafferty (Ephemeroptera, Baetidae), with description of a new subgenus and 20 new species. ZooKeys 1227: 159-347, DOI: 10.3897/zookeys.1227.138100
835CD3DA6248573393280208AFAD25D1.taxon	distribution	Distribution. New Guinea (Fig. 146).	en	Kaltenbach, Thomas, Kluge, Nikita J., Gattolliat, Jean-Luc (2025): Redescription of Papuanatula Lugo-Ortiz & McCafferty (Ephemeroptera, Baetidae), with description of a new subgenus and 20 new species. ZooKeys 1227: 159-347, DOI: 10.3897/zookeys.1227.138100
77FD5C6C0E195936AA811122427BE6EA.taxon	description	Figs 38, 39, 40, 41, 42, 43	en	Kaltenbach, Thomas, Kluge, Nikita J., Gattolliat, Jean-Luc (2025): Redescription of Papuanatula Lugo-Ortiz & McCafferty (Ephemeroptera, Baetidae), with description of a new subgenus and 20 new species. ZooKeys 1227: 159-347, DOI: 10.3897/zookeys.1227.138100
77FD5C6C0E195936AA811122427BE6EA.taxon	etymology	Etymology. The species name is based on the Latin words dum spinae meaning “ long spines ”, referring to the long denticles at posterior margins of abdominal terga.	en	Kaltenbach, Thomas, Kluge, Nikita J., Gattolliat, Jean-Luc (2025): Redescription of Papuanatula Lugo-Ortiz & McCafferty (Ephemeroptera, Baetidae), with description of a new subgenus and 20 new species. ZooKeys 1227: 159-347, DOI: 10.3897/zookeys.1227.138100
77FD5C6C0E195936AA811122427BE6EA.taxon	diagnosis	Diagnosis. Larva. The following combination of characters distinguishes P. dumspinae sp. nov. from other species of Papuanatula s. str.: body dorsally with row of long, fine, simple setae along midline; abdominal terga without protuberances; femur basally with wedge-shaped blank, medial area dark grey, distal area yellow-brown; paracercus with eight or nine segments; abdominal terga with very long, narrow, triangular denticles on posterior margins.	en	Kaltenbach, Thomas, Kluge, Nikita J., Gattolliat, Jean-Luc (2025): Redescription of Papuanatula Lugo-Ortiz & McCafferty (Ephemeroptera, Baetidae), with description of a new subgenus and 20 new species. ZooKeys 1227: 159-347, DOI: 10.3897/zookeys.1227.138100
77FD5C6C0E195936AA811122427BE6EA.taxon	description	Description. Larva (Figs 38 – 43). Body length 3.8 – 5.2 mm, cerci much longer than body length (~ 1.5 ×). Cuticular coloration (Fig. 38 a – c). Head, thorax and abdomen dorsally brown; thorax with complex pattern; abdominal segments II – IV and VII – IX dark brown to black, V, VI and X yellow-brown. Femur basally with wedge-shaped blank, medial area dark grey, distal area yellow-brown; tibia ecru with grey; tarsus and claw grey-brown. Thorax and abdominal segment I ventrally ecru, protuberances of thoracic sterna dark brown, abdominal segments II – X ventrally brown. Cerci grey-brown. Hypodermal coloration. Each abdominal tergum I – IX with wide dark brown transverse band close to anterior margin and with narrower dark brown transverse band close to posterior margin (Fig. 38 a). Head. Dorsally with irregular row of long, fine, simple setae along midline. Antenna (Fig. 41 i). Length ~ 1.5 × head length. As typical for subgenus. Developing turbinate eyes in mature male larva (Fig. 41 i) ovoid, with large distance to each other. Labrum (Fig. 39 a, b). Length 0.5 × maximum width, laterally convex. Dorsal, sub-marginal arc with 22 – 29 feathered setae. Right mandible (Fig. 39 d, e). Margin between prostheca and mola with row of minute denticles. Otherwise, as typical for subgenus. Left mandible (Fig. 39 f, g). Margin between prostheca and mola with row of minute denticles. Otherwise, as typical for subgenus. Hypopharynx (Fig. 39 c). As typical for genus. Maxilla (Fig. 40 c, d). Maxillary palp somewhat shorter than galea-lacinia, robust; palp segment II ~ 1.3 × length of segment I. Otherwise, as typical for genus. Labium (Fig. 40 a, b). As typical for the genus. Paraglossa dorsally with two spine-like setae near inner, distolateral margin. Labial palp with segment I ~ 1.1 × length of segments II and III combined. Segment II with minute distomedial protuberance, dorsally with row of four spine-like setae near outer, distolateral margin. Segment III slightly pentagonal, pointed; 0.7 × length of segment II; inner dorsal margin with few feathered setae. Thorax. Sterna. With small protuberances on sides of prosternum and close to openings of mesothoracic and metathoracic sternal apodemes (as in Fig. 108 a). Terga (Fig. 38 b) without protuberances; with row of long, fine, simple setae along midline. Legs (Fig. 41 a – h). Ratio of leg segments: fore leg 0.9: 1.0: 0.3: 0.1, middle leg 0.9: 1.0: 0.3: 0.1 and hind leg 1.1: 1.0: 0.3: 0.2. Femur. Length ~ 3.5 × maximum width. Claw with one row of 6 – 8 denticles and one posterior seta. Abdomen. Terga (Figs 42 a, 43 a – c) with row of long, fine, simple setae along midline. Terga without protuberances, terga I – IV with slight, paired, medioposterior elevations. Posterior margin of terga: (I) II – IX with long, narrow, triangular, pointed denticles. Surface with scattered small, paddle-like, striated scales with slightly serrate margin. Tergalii (Fig. 42 e, f). Broad ovoid, tracheation well developed; margins smooth, with few short, fine, simple setae. Paraproct (Fig. 42 g, h). Posterior margin with prolongation and row of minute denticles; on surface an area with minute, notched scales. Caudalii (Fig. 42 b – d). Cerci apart from basal and distal part on ¾ of their length with up to ten swimming setae per segment, initially increasing and then again decreasing toward distal part. Paracercus with eight or nine segments. Pose of subimaginal gonostyli under larval cuticle. Unknown. Subimago. Unknown. Imago. Unknown. Egg. Unknown.	en	Kaltenbach, Thomas, Kluge, Nikita J., Gattolliat, Jean-Luc (2025): Redescription of Papuanatula Lugo-Ortiz & McCafferty (Ephemeroptera, Baetidae), with description of a new subgenus and 20 new species. ZooKeys 1227: 159-347, DOI: 10.3897/zookeys.1227.138100
77FD5C6C0E195936AA811122427BE6EA.taxon	distribution	Distribution. New Guinea (Fig. 146).	en	Kaltenbach, Thomas, Kluge, Nikita J., Gattolliat, Jean-Luc (2025): Redescription of Papuanatula Lugo-Ortiz & McCafferty (Ephemeroptera, Baetidae), with description of a new subgenus and 20 new species. ZooKeys 1227: 159-347, DOI: 10.3897/zookeys.1227.138100
499A62E183BC5CA0A698FC1A4219FE9E.taxon	description	Figs 44, 45, 46, 47	en	Kaltenbach, Thomas, Kluge, Nikita J., Gattolliat, Jean-Luc (2025): Redescription of Papuanatula Lugo-Ortiz & McCafferty (Ephemeroptera, Baetidae), with description of a new subgenus and 20 new species. ZooKeys 1227: 159-347, DOI: 10.3897/zookeys.1227.138100
499A62E183BC5CA0A698FC1A4219FE9E.taxon	etymology	Etymology. The species name duplex is based on the Latin word for “ double ”, referring to the paired protuberances on abdominal terga.	en	Kaltenbach, Thomas, Kluge, Nikita J., Gattolliat, Jean-Luc (2025): Redescription of Papuanatula Lugo-Ortiz & McCafferty (Ephemeroptera, Baetidae), with description of a new subgenus and 20 new species. ZooKeys 1227: 159-347, DOI: 10.3897/zookeys.1227.138100
499A62E183BC5CA0A698FC1A4219FE9E.taxon	diagnosis	Diagnosis. Larva. The following combination of characters distinguishes P. duplex sp. nov. from other species of Papuanatula s. str.: body dorsally without row of long, fine, simple setae along midline; pronotum dorsally with posteromedial, paired, triangular protuberances; metanotum dorsally with posteromedial, paired, long, cylindrical, distally slightly conical protuberances; abdomen dorsally with posteromedial, paired protuberances: terga I – VI long, subcylindrical, distally slightly conical; terga VII – IX shorter, compressed, triangular.	en	Kaltenbach, Thomas, Kluge, Nikita J., Gattolliat, Jean-Luc (2025): Redescription of Papuanatula Lugo-Ortiz & McCafferty (Ephemeroptera, Baetidae), with description of a new subgenus and 20 new species. ZooKeys 1227: 159-347, DOI: 10.3897/zookeys.1227.138100
499A62E183BC5CA0A698FC1A4219FE9E.taxon	description	Description. Larva (Figs 44 – 47). Body length 4.5 mm, cerci broken. Cuticular coloration (Fig. 44 a – c). Head, thorax and abdomen dorsally yellow-brown to brown; metanotum and abdominal terga I – IV darker than V – IX; Head, thorax, and abdominal segment I ventrally pale yellow-brown, abdominal segments II – IX yellow-brown; posterolateral protuberances on thorax brown. Legs yellow-brown; femur with long, narrow blank along dorsal margin. Hypodermal coloration (Fig. 44 a). Abdominal segments I – IX dorsally with transverse band along posterior margins. Head. Antenna (Fig. 44 a, b). Length ~ 1.5 × head length. Developing turbinate eyes in last instar male larva unknown. Labrum (Fig. 45 a, b). Length 0.5 × maximum width, laterally convex. Dorsal, sub-marginal arc with ~ 12 feathered setae. Right mandible (Fig. 45 d, e). Margin between prostheca and mola with row of short denticles. Otherwise, as typical for subgenus. Left mandible (Fig. 45 f, g). Margin between prostheca and mola with row of short denticles. Otherwise, as typical for subgenus. Hypopharynx (Fig. 45 c). As typical for genus. Maxilla (Fig. 45 j, k). Maxillary palp subequal in length to galea-lacinia; palp segment II slightly longer than segment I. Otherwise, as typical for genus. Labium (Fig. 45 h, i). As typical for the genus. Paraglossa with two spine-like setae on inner, distolateral margin. Labial palp with segment I 0.9 × length of segments II and III combined. Segment II without distomedial protuberance, dorsally with row of five spine-like setae near outer, distolateral margin. Segment III slightly oblong, pointed, as long as segment II. Thorax. Sterna. With small protuberances on sides of prosternum and close to openings of mesothoracic and metathoracic sternal apodemes (as in Fig. 108 a). Terga (Figs 44 a, b, 47 a – c) Pronotum with posteromedial, paired, triangular protuberances; metanotum with posteromedial, paired, long, cylindrical, distally slightly conical protuberances. Legs (Fig. 46 a – c). Ratio of leg segments: fore leg 1.0: 1.0: 0.3: 0.2, middle leg 1.0: 1.0: 0.3: 0.2 and hind leg 1.0: 1.0: 0.4: 0.2. Femur. Length ~ 3 × maximum width. Claw with one row of eight denticles; one posterior seta. Abdomen. Terga (Figs 44 a, b, 46 d, e, 47 a, b, d) I – IX with posteromedial, paired protuberances: I – VI long, subcylindrical, distally slightly conical; VII – IX shorter, compressed, triangular. Posterior margin of terga: I – V unknown; VI – IX with minute denticles, apically split with several points. Surface with scattered small, triangular, pointed, striated scales. Tergalii (Fig. 46 g) ovoid, tracheation developed; margins smooth, with few short, fine, simple setae. Paraproct (Fig. 46 f). Posterior margin with prolongation and row of minute denticles. Caudalii. Unknown. Pose of subimaginal gonostyli under larval cuticle. Unknown. Subimago. Unknown. Imago. Unknown. Egg. Unknown.	en	Kaltenbach, Thomas, Kluge, Nikita J., Gattolliat, Jean-Luc (2025): Redescription of Papuanatula Lugo-Ortiz & McCafferty (Ephemeroptera, Baetidae), with description of a new subgenus and 20 new species. ZooKeys 1227: 159-347, DOI: 10.3897/zookeys.1227.138100
499A62E183BC5CA0A698FC1A4219FE9E.taxon	distribution	Distribution. New Guinea (Fig. 146).	en	Kaltenbach, Thomas, Kluge, Nikita J., Gattolliat, Jean-Luc (2025): Redescription of Papuanatula Lugo-Ortiz & McCafferty (Ephemeroptera, Baetidae), with description of a new subgenus and 20 new species. ZooKeys 1227: 159-347, DOI: 10.3897/zookeys.1227.138100
106D5D67C34850D18EA366C9811F1CF0.taxon	description	Figs 48, 49, 50, 51, 52, 53, 54	en	Kaltenbach, Thomas, Kluge, Nikita J., Gattolliat, Jean-Luc (2025): Redescription of Papuanatula Lugo-Ortiz & McCafferty (Ephemeroptera, Baetidae), with description of a new subgenus and 20 new species. ZooKeys 1227: 159-347, DOI: 10.3897/zookeys.1227.138100
106D5D67C34850D18EA366C9811F1CF0.taxon	etymology	Etymology. The species name epibessa refers to the morphological similarity with P. bessa.	en	Kaltenbach, Thomas, Kluge, Nikita J., Gattolliat, Jean-Luc (2025): Redescription of Papuanatula Lugo-Ortiz & McCafferty (Ephemeroptera, Baetidae), with description of a new subgenus and 20 new species. ZooKeys 1227: 159-347, DOI: 10.3897/zookeys.1227.138100
106D5D67C34850D18EA366C9811F1CF0.taxon	diagnosis	Diagnosis. Larva. The following combination of characters distinguishes P. epibessa sp. nov. from other species of Papuanatula s. str.: body dorsally with irregular row of long, fine, simple setae along midline; metanotum and abdominal terga I – III with medioposterior, broad, paired humps, poorly developed on terga I – III; femur yellow brown, basally with wedge-shaped blank; paracercus with 6 – 8 segments; claw with 1 – 3 posterior setae.	en	Kaltenbach, Thomas, Kluge, Nikita J., Gattolliat, Jean-Luc (2025): Redescription of Papuanatula Lugo-Ortiz & McCafferty (Ephemeroptera, Baetidae), with description of a new subgenus and 20 new species. ZooKeys 1227: 159-347, DOI: 10.3897/zookeys.1227.138100
106D5D67C34850D18EA366C9811F1CF0.taxon	description	Description. Larva (Figs 48 – 54). Body length 4.1 – 5.5 mm, cerci much longer than body length (~ 1.7 ×). Cuticular coloration (Figs 48 a – c, 49 a) Head, thorax and abdomen dorsally yellow brown to brown, thorax with complex markings. Abdominal terga II – V with brown markings along anterior margin, tergum III additionally with mediolateral, brown markings. Femur yellow brown, basally with wedge-shaped blank. Tibia pale yellow brown; tarsus yellow brown. Cerci pale yellow brown. Hypodermal coloration (Figs 48 a, 49 a) Abdominal terga with narrow, dark brown, transverse bands along posterior margin. Head. Dorsally with irregular row of long, fine, simple setae along midline. Antenna (Fig. 51 e). Length ~ 1.5 × head length. Developing turbinate eyes in last instar male larva (Fig. 51 e) rather small, ovoid, with big distance to each other. Labrum (Fig. 50 a, b). Length 0.5 × maximum width, laterally convex. Dorsal, sub-marginal arc with ~ 35 feathered setae. Right mandible (Fig. 50 d, e). Margin between prostheca and mola with few denticles close to prostheca. Otherwise, as typical for subgenus. Left mandible (Fig. 50 f, g). Margin between prostheca and mola smooth, with few denticles close to prostheca. Otherwise, as typical for subgenus. Hypopharynx (Fig. 50 c). As typical for genus. Maxilla (Fig. 51 c, d). Maxillary palp subequal in length to galea-lacinia; palp segment II approximately as long as segment I. Otherwise, as typical for genus. Labium (Fig. 51 a, b). As typical for the genus. Paraglossa dorsally with 2 spine-like setae near inner, distolateral margin. Labial palp with segment I approximately as long as segments II and III combined. Segment II with minute distomedial protuberance, dorsally with row of five or six spine-like setae near outer, distolateral margin. Segment III conical, pointed; 0.8 × length of segment II; inner dorsal margin with few feathered setae. Thorax. Sterna. With small protuberances on sides of prosternum and close to openings of mesothoracic and metathoracic sternal apodemes (as Fig. 108 a). Terga (Figs 48 b, 53 a) with irregular row of long, fine, simple setae along midline. Metanotum with medioposterior broad, paired humps, without hind protoptera or their vestiges. Legs (Fig. 52 a – h). Ratio of leg segments: fore leg 0.8: 1.0: 0.3: 0.1, middle leg 0.9: 1.0: 0.3: 0.1 and hind leg 1.0: 1.0: 0.3: 0.1. Femur. Length ~ 4 × maximum width. Posterior side of apex with row of robust setae (contrary to fine, simple setae as it is usually the case in Papuanatula s. str.). Tarsus. Inner margin distally sometimes with two long setae (Fig. 52 g). Claw with one row of 6 – 8 (9) denticles and 1 – 3 posterior setae. Abdomen. Terga (Figs 48 b, 53 a, b, 54 a, b) with irregular row of long, fine, simple setae along midline. Terga I – III with poorly developed, medioposterior, broad, paired humps. Terga II – IV protruding slightly medially on posterior margin. Posterior margin of terga: I smooth, without denticles; II – IX with triangular, apically rounded denticles, partly with minute, pointed denticles in between. Surface with scattered small, ovoid, striated scales with slightly serrate margin. Tergalii (Fig. 53 e, f). Ovoid, tracheation poorly developed or absent; margins smooth, with short, fine, simple setae. Paraproct (Fig. 53 g). Posterior margin with prolongation and row of minute denticles. Caudalii (Fig. 53 c, d). Cerci in middle part with one to maximally five swimming setae per segment, initially increasing and then again decreasing toward distal part; sometimes total loss or maximally one swimming seta per segment. Paracercus with 6 – 8 segments. Pose of subimaginal gonostyli under larval cuticle. Unknown. Subimago. Unknown. Imago. Unknown. Egg. Unknown.	en	Kaltenbach, Thomas, Kluge, Nikita J., Gattolliat, Jean-Luc (2025): Redescription of Papuanatula Lugo-Ortiz & McCafferty (Ephemeroptera, Baetidae), with description of a new subgenus and 20 new species. ZooKeys 1227: 159-347, DOI: 10.3897/zookeys.1227.138100
106D5D67C34850D18EA366C9811F1CF0.taxon	distribution	Distribution. New Guinea (Fig. 146).	en	Kaltenbach, Thomas, Kluge, Nikita J., Gattolliat, Jean-Luc (2025): Redescription of Papuanatula Lugo-Ortiz & McCafferty (Ephemeroptera, Baetidae), with description of a new subgenus and 20 new species. ZooKeys 1227: 159-347, DOI: 10.3897/zookeys.1227.138100
0F1C9FB320815E0980C3B0BE679108BF.taxon	description	Figs 55, 56, 57, 58, 59, 60, 61	en	Kaltenbach, Thomas, Kluge, Nikita J., Gattolliat, Jean-Luc (2025): Redescription of Papuanatula Lugo-Ortiz & McCafferty (Ephemeroptera, Baetidae), with description of a new subgenus and 20 new species. ZooKeys 1227: 159-347, DOI: 10.3897/zookeys.1227.138100
0F1C9FB320815E0980C3B0BE679108BF.taxon	etymology	Etymology. The species name heterochaeta refers to the sharp difference between blunt-ended setae which form the longitudinal row on outer side of larval femur and small hair-like setae which form continuation of this row on apex of the femur (Fig. 58 a).	en	Kaltenbach, Thomas, Kluge, Nikita J., Gattolliat, Jean-Luc (2025): Redescription of Papuanatula Lugo-Ortiz & McCafferty (Ephemeroptera, Baetidae), with description of a new subgenus and 20 new species. ZooKeys 1227: 159-347, DOI: 10.3897/zookeys.1227.138100
0F1C9FB320815E0980C3B0BE679108BF.taxon	diagnosis	Diagnosis. Larva. The following combination of characters distinguishes P. heterochaeta sp. nov. from other species of Papuanatula s. str.: body dorsally without irregular row of long, fine, simple setae along midline; abdominal terga without protuberances; hypopharynx apically with pair of bunches of setae-like spines (instead of one bunch as usually); femur with large, proximal, oval blank and with dark brown, shoe-shaped macula inside proximal blank; sharp difference between setae which form the longitudinal row on outer side of femur and setae which form continuation of this row on apex of femur; long, slender, preapical seta on tarsus absent; paracercus with ~ 10 – 12 segments.	en	Kaltenbach, Thomas, Kluge, Nikita J., Gattolliat, Jean-Luc (2025): Redescription of Papuanatula Lugo-Ortiz & McCafferty (Ephemeroptera, Baetidae), with description of a new subgenus and 20 new species. ZooKeys 1227: 159-347, DOI: 10.3897/zookeys.1227.138100
0F1C9FB320815E0980C3B0BE679108BF.taxon	description	Description. Larva (Figs 55 – 59). Cuticular coloration. Head, pronotum, mesonotum and metanotum brownish with paler areas; fore protoptera with narrow paler lines corresponding to some longitudinal veins (Fig. 56 a, b). Thoracic pleura brownish, sterna mostly colorless. Cuticle of femur with brownish margins and brownish transverse band separating large proximal blank from smaller distal blank; proximal blank oval (not wedge-shaped), occupying most part of proximal 1 / 2 (Fig. 56 d – f). Tibia and tarsus mostly brownish (Fig. 56 d – f). Abdominal terga brownish with paler blanks; median blank on tergum VI larger than others. Sterna mostly colorless (Fig. 56 c). Cerci uniformly pale brownish. Hypodermal coloration. Anterior side of each femur with dark brown shoe-shaped macula on proximal 1 / 2, within proximal cuticular blank (Fig. 55 f, g); posterior side of each femur with or without two longitudinal, brown maculae (Fig. 55 d), sometimes connected one with another (Fig. 55 b). Boundaries between abdominal terga narrowly bordered by dark brown; other brown markings on abdomen absent or present, most extensive on abdominal tergum IV (Fig. 55 a). Tissues surrounding tracheae of tergalii (main trachea and its branches) with brown pigmentation (Fig. 59 d – i). Head. Antenna. As typical for subgenus. Developing turbinate eyes in last instar male larva with facets equally developed on middle and peripheric areas (as in Fig. 32 d). Labrum (Fig. 57 a, b) very slightly widened distally; long setae on dorsal surface spaced and forming regular transverse row; each seta pointed, with moderately long processes on both sides. Right mandible (Fig. 57 d, f). As typical for subgenus. Left mandible (Fig. 57 c, e). Incisor and kinetodontium non-distinguishable, together with 5 denticles proximad of stretched apex of incisor. Otherwise, as typical for subgenus. Hypopharynx (Fig. 57 j) apically with pair of bunches of stout setae-like spines. Maxilla (Fig. 57 g). Maxillary palps long as galea-lacinia. Otherwise, as typical for genus. Labium (Fig. 57 h, i). Paraglossae widest at base and narrowing toward apex; three apical setal rows bent at apex of paraglossa. Glossa shorter than half of paraglossa, with finger-like (distal) portion as long as triangular (proximal) portion. Glossa with several long setae at apex and one long seta near middle of ventral side. Labial palp without distomedian projection on segment II; segment III with median margin as long as lateral margin. Thorax. Sterna without protuberances. Terga. Without long setae along midline. Metanotum without hind protoptera or their vestiges. Legs (Figs 56 d – f, 58 a – f). Fore femur widened in proximal part; hind tibia shorter than others. Femur. Outer side of each femur with single regular row of long setae; each seta slender, flattened, narrowing toward apex and blunt apically, with numerous fine, short branches on all sides. Distally, close to femur-tibia articulation, setal row continued by several smaller, hair-like setae (similar to setae on tibia). Anterior side of femur with small, stout setae, sparsely and irregularly situated. Serrate area located at middle of anterior side, partly on brown transverse band, partly on proximal blank. Tibia. Patella-tibial suture present on all legs, terminated near middle of inner margin of tibia. Tibia-tarsal condylus turned to anterior side. Anterior side of each tibia with regular row of hair-like setae similar to setae near apex of femur. Tarsus. Anterior side of each tarsus with regular row of similar, but shorter (not narrower) setae. Long preapical seta absent; posterior side of each tarsus with regular row of few very short, stout setae and one longer, pointed seta of same thickness distad of them (Fig. 58 b, c). Claw with row of 6 – 9 denticles and one somewhat larger denticle distad of them; long, arched, posterior seta (Fig. 58 e, f). Abdomen. Terga (Figs 56 c, 59 a, b) without dorsal protuberances, only with slightly expressed, unpaired, median elevations; without long setae along midline. Abdominal terga with numerous small scales with small sockets and fan-like striation. Abdominal terga I – III without denticles on posterior margins; posterior margins of abdominal terga IV – IX with small, pointed denticles. Posterior margin of tergum X with very small denticles. Tergalii (Fig. 59 d – i) of abdominal segment I absent; tergalii II – VII subequal, oval, relatively narrow. Each tergalius with costal and anal ribs narrow, smooth, present on proximal 1 / 2 of tergalius only. Paraproct (Fig. 59 c). Margins membranous and smooth, lacking denticles. Caudalii (Fig. 59 c) without swimming setae or their vestiges. Paracercus short, consisting of ~ 10 – 12 segments. Pose of subimaginal gonostyli under larval cuticle (Fig. 61 a). In mature larva ready to molt to subimago, subimaginal gonostyli packed under larval cuticle in “ Labiobaetis - type ” pose, as typical for the genus. 2 nd segment directed medially and bent proximally; 3 rd segment directed medially (as continuation of 2 nd segment) and narrowed apically, being deformed corresponding to space between subimaginal styliger and larval cuticle. Subimago. Cuticular coloration. Pronotum and prosternum partly brown (Fig. 60 f). Mesonotum pale brown with medioparapsidal suture colorless, other sutures darker brown (Fig. 60 e). Meso- and metathoracic pleura and sterna with colorless, pale brownish and dark brown areas (Fig. 60 f). Cuticle of wings colorless, with microtrichiae brownish. Legs nearly colorless, with pale brown bordering on femur and base of tibia (Fig. 60 d). Abdomen very pale brownish with colorless sigilla. Cerci colorless with setae brownish. Hypodermal coloration. As in imago. Texture. On all legs of both sexes, each tarsomere covered mostly with blunt microlepides, with pointed microlepides near apex (as in Fig. 70 i). Imago. Imago, male. Head ochre. Antennae ochre. Turbinate eyes red, widened apically. Thorax ochre, equally pale dorsally, laterally, and ventrally, with dark brown hypodermal markings on lateral sides. Fore wing with membrane colorless, veins ochre. Pterostigma with three or four incomplete, oblique cross veins (Fig. 60 a – c). Legs mostly ochre; on leg of each pair, anterior side of femur with contrasting, dark brown, longitudinal macula just proximad of midlength; posterior side of femur with reddish brown macula near apex (Fig. 60 g). Abdomen mostly whitish or ochre; each tergum I – IX with dark brown transverse band close to posterior margin; each tergum III and IV with pair of brown spots; tergum IV with transverse brown macula posteriad-mediad of them (Fig. 60 a – c). Cerci ochre. Genitalia (Fig. 61 c, d). Sterno-styligeral muscle absent. Each unistyliger sharply widened apically on median side, so that median margins convergent distally. 1 st segment of gonostylus with lateral side convex, median side median side gradually turns to 2 nd segment. Second segment equally wide all over its length. Third (terminal) segment of gonostylus nearly as wide as 2 nd, with length slightly exceeding width. Penial bridge with wide, blunt, membranous projection between unistyligers. Each gonovectis parabolic, with lateral (basal) and median (apical) portions equally long, apex bent medially-caudally. Imago, female (Fig. 60 d). Unknown. Judging by subimago, coloration of head, thorax, and legs similar to that of male; abdominal terga with ochre-brown pigmentation. Egg. Unknown.	en	Kaltenbach, Thomas, Kluge, Nikita J., Gattolliat, Jean-Luc (2025): Redescription of Papuanatula Lugo-Ortiz & McCafferty (Ephemeroptera, Baetidae), with description of a new subgenus and 20 new species. ZooKeys 1227: 159-347, DOI: 10.3897/zookeys.1227.138100
0F1C9FB320815E0980C3B0BE679108BF.taxon	distribution	Distribution. New Guinea (Fig. 147).	en	Kaltenbach, Thomas, Kluge, Nikita J., Gattolliat, Jean-Luc (2025): Redescription of Papuanatula Lugo-Ortiz & McCafferty (Ephemeroptera, Baetidae), with description of a new subgenus and 20 new species. ZooKeys 1227: 159-347, DOI: 10.3897/zookeys.1227.138100
F7F3649175985A59BCB1B831EB479A93.taxon	type_taxon	Type species. Papuanatula (Papuanatula) tuber Lugo-Ortiz & McCafferty, 1999.	en	Kaltenbach, Thomas, Kluge, Nikita J., Gattolliat, Jean-Luc (2025): Redescription of Papuanatula Lugo-Ortiz & McCafferty (Ephemeroptera, Baetidae), with description of a new subgenus and 20 new species. ZooKeys 1227: 159-347, DOI: 10.3897/zookeys.1227.138100
F7F3649175985A59BCB1B831EB479A93.taxon	diagnosis	Diagnosis (larval characters). Distal part of antenna asymmetrical: each flagellomere with anterior side more convex than posterior side, with brown hypodermal spot near apex of anterior side (Fig. 2 b). Labrum dorsally with submarginal row of long, feathered setae (Fig. 3 b). Each seta consists of a stout stem and long processes on both sides. Mandibles. Both mandibles long, with incisor strongly elongated (in contrast to shortened in Papuafiliola nov.); full-length mandibles present only at the beginning of each instar, often worn at the end of instar. Right mandible (Fig. 3 e): kinetodontium deeply separated from incisor; incisor with three denticles; kinetodontium terminating with three or four denticles, with distal denticle longest. Left mandible (Fig. 3 f): incisor and kinetodontium fused at most of its length; incisor with three denticles; kinetodontium terminating with three or four denticles, with distal denticle longest. Labium: glossa usually approximately as long as half of paraglossa; with several long setae at apex and usually one long seta near middle of ventral side. Labial palp without real distomedian projection on segment II (Fig. 3 c) (in contrast to Papuafiliola subgen. nov.). Legs (Figs 4 a – e, 108 b): Outer side of each femur usually with single regular row of long, hair-like setae bearing numerous fine, short branches on all sides. Anterior side of each tibia usually with regular row of setae similar to that on femur. Anterior side of each tarsus usually with regular row of setae similar to femur and tibia. Posterior side of each tarsus with regular row of short, stout setae and one (rarely two) much longer, thinner, pointed seta distad. Paraproct (Fig. 12 e): posterior margin usually with prolongation. Pose of subimaginal gonostyli under larval cuticle (Fig. 76 b). Subimaginal gonostyli developing under cuticle of last instar male larvae folded in “ Labiobaetis - type ” (Kluge 2004: fig. 29 I): 1 st segment directed laterally, 2 nd and 3 rd segments directed medially, whereby the 2 nd segment is the closest to the posterior margin of the 9 th segment. Eggs of examined species without regular relief (Fig. 37 d, e).	en	Kaltenbach, Thomas, Kluge, Nikita J., Gattolliat, Jean-Luc (2025): Redescription of Papuanatula Lugo-Ortiz & McCafferty (Ephemeroptera, Baetidae), with description of a new subgenus and 20 new species. ZooKeys 1227: 159-347, DOI: 10.3897/zookeys.1227.138100
1621D2C3529B5AFD9CA65A062B5BA7FE.taxon	description	Figs 14, 15, 16	en	Kaltenbach, Thomas, Kluge, Nikita J., Gattolliat, Jean-Luc (2025): Redescription of Papuanatula Lugo-Ortiz & McCafferty (Ephemeroptera, Baetidae), with description of a new subgenus and 20 new species. ZooKeys 1227: 159-347, DOI: 10.3897/zookeys.1227.138100
1621D2C3529B5AFD9CA65A062B5BA7FE.taxon	diagnosis	Diagnosis. Larva. The following combination of characters distinguishes P. lenos from other species of Papuanatula s. str.: body dorsally without row of long, fine, simple setae along midline; body dorsally without protuberances; thorax ventrally without protuberances; thorax dorsally without distinct markings; femur anteriorly with hypodermal, large, oblong to drop-shaped, dark brown to purple black marking in basal 1 / 2; paracercus with nine segments (immature larvae 11); paraproct without posterior prolongation;	en	Kaltenbach, Thomas, Kluge, Nikita J., Gattolliat, Jean-Luc (2025): Redescription of Papuanatula Lugo-Ortiz & McCafferty (Ephemeroptera, Baetidae), with description of a new subgenus and 20 new species. ZooKeys 1227: 159-347, DOI: 10.3897/zookeys.1227.138100
1621D2C3529B5AFD9CA65A062B5BA7FE.taxon	description	Description. Larva (Figs 14 – 16; Lugo-Ortiz and McCafferty 1999: 65 – 66, figs 19 – 24). Body length 3.3 – 4.4 mm, cerci length unknown. Cuticular coloration (Fig. 14 a). Head, thorax and abdomen dorsally yellow-brown to brown with pattern as in Fig. 14 a, slightly variable; abdominal segments III – VIII with oblique, dark brown lateral markings, partly forming a trough-shaped pattern (segments III, VII, and VIII), segment IV with dark brown crown-like pattern. Legs yellow-brown to brown; Caudalii yellow-brown. Hypodermal coloration. Abdominal segments I – IX dorsally with transverse stripe on posterior margins (Fig. 14 a). Femur anteriorly with large oblong to drop-shaped, dark brown to purple black marking in basal 1 / 2; posteriorly with long, broad, dark brown to purple black dashes. Head. Antenna. Length 1.5 × head length. Developing turbinate eyes in last instar male larva unknown. Labrum (Fig. 15 a). Length 0.5 × maximum width, laterally convex. Dorsal, sub-marginal arc with 12 – 15 feathered setae. Right mandible (Fig. 15 c, d). Margin between prostheca and mola smooth. Otherwise, as typical for subgenus. Left mandible (Fig. 15 e, f). Margin between prostheca and mola smooth, with one spine close to subtriangular process. Otherwise, as typical for subgenus. Hypopharynx (Fig. 15 b). As typical for genus. Maxilla (Fig. 15 h, i). Maxillary palp subequal in length to galea-lacinia; palp segment II subequal in length to segment I. Otherwise, as typical for genus. Labium (Fig. 15 g). As typical for genus. Paraglossa dorsally with one spine-like seta near inner, distolateral margin. Labial palp with segment I subequal in length to segments II and III combined. Segment II without distomedial protuberance, dorsally with row of five spine-like setae near outer, distolateral margin. Segment III bulbous, pointed, 0.8 × length of segment II; inner dorsal margin with few feathered setae. Thorax. Sterna without protuberances. Terga without protuberances. Legs (Fig. 16 a, b). Ratio of leg segments: fore leg 1.0: 1.0: 0.2: 0.1, middle leg 0.9: 1.0: 0.2: 0.1 and hind leg 1.1: 1.0: 0.3: 0.1. Femur. Length ~ 3 × maximum width. Claw with one row of six or seven denticles, one posterior seta. Abdomen. Terga (Fig. 16 c). Abdominal terga without protuberances. Posterior margin of terga: I smooth, without denticles; II – IX with triangular, pointed denticles. Tergalii (Fig. 16 e – f) ovoid, tracheation rather poorly developed; margins smooth, with short, fine, simple setae. Paraproct. Posterior margin without prolongation; smooth, without denticles. Caudalii (Fig. 16 d). Paracercus with nine segments, immature larvae up to 11 segments. Pose of subimaginal gonostyli under larval cuticle. Unknown. Subimago. Unknown. Imago. Unknown. Egg. Unknown.	en	Kaltenbach, Thomas, Kluge, Nikita J., Gattolliat, Jean-Luc (2025): Redescription of Papuanatula Lugo-Ortiz & McCafferty (Ephemeroptera, Baetidae), with description of a new subgenus and 20 new species. ZooKeys 1227: 159-347, DOI: 10.3897/zookeys.1227.138100
1621D2C3529B5AFD9CA65A062B5BA7FE.taxon	distribution	Distribution. New Guinea (Fig. 146).	en	Kaltenbach, Thomas, Kluge, Nikita J., Gattolliat, Jean-Luc (2025): Redescription of Papuanatula Lugo-Ortiz & McCafferty (Ephemeroptera, Baetidae), with description of a new subgenus and 20 new species. ZooKeys 1227: 159-347, DOI: 10.3897/zookeys.1227.138100
44836D371795592F96323CCBE3EAB69A.taxon	description	Figs 62, 63, 64	en	Kaltenbach, Thomas, Kluge, Nikita J., Gattolliat, Jean-Luc (2025): Redescription of Papuanatula Lugo-Ortiz & McCafferty (Ephemeroptera, Baetidae), with description of a new subgenus and 20 new species. ZooKeys 1227: 159-347, DOI: 10.3897/zookeys.1227.138100
44836D371795592F96323CCBE3EAB69A.taxon	etymology	Etymology. The species name normungulata refers to the larval claw structure, which lacks the arched posterior seta and the enlarged denticle associated with this seta that is normal for the plesiomorphon Papuanatula, being different from other species of Papuanatula.	en	Kaltenbach, Thomas, Kluge, Nikita J., Gattolliat, Jean-Luc (2025): Redescription of Papuanatula Lugo-Ortiz & McCafferty (Ephemeroptera, Baetidae), with description of a new subgenus and 20 new species. ZooKeys 1227: 159-347, DOI: 10.3897/zookeys.1227.138100
44836D371795592F96323CCBE3EAB69A.taxon	diagnosis	Diagnosis. Larva. The following combination of characters distinguishes P. normungulata sp. nov. from other species of Papuanatula s. str.: body without irregular row of long, fine, simple setae along midline; abdominal terga I – VIII with unpaired, long, pointed protuberance close to posterior margin; patella-tibial suture reduced; posterior seta absent; wide stripe of densely situated setae instead of the regular setal row on outer side of femur and tibia.	en	Kaltenbach, Thomas, Kluge, Nikita J., Gattolliat, Jean-Luc (2025): Redescription of Papuanatula Lugo-Ortiz & McCafferty (Ephemeroptera, Baetidae), with description of a new subgenus and 20 new species. ZooKeys 1227: 159-347, DOI: 10.3897/zookeys.1227.138100
44836D371795592F96323CCBE3EAB69A.taxon	description	Description. Larva (Figs 62 – 64). Cuticular coloration. Head brownish. Pronotum and mesonotum pale brownish with darker areas; fore protoptera with wide darker lines corresponding to convex veins and thin paler lines corresponding to concave veins (Figs 62 a, 63 a, b). Cuticle of femur mostly pale ochre-brownish, darker brown at apex. Tibia and tarsus ochre-brownish (Fig. 64 a, b). Abdominal terga ochre-brownish with median spines darker brown. Cerci brow. Hypodermal coloration. In female larva with developed subimaginal wings, whole dorsal side of head, thorax, and abdomen uniformly dark brown; pleura of thorax ochre with dark brown; prosternum and mesosternum dark brown; abdominal sterna with ochre and brown areas (Figs 62 a, 63 a, b). On anterior side of each leg, proximal ¾ of femur entirely dark brown, distal ¼ ochre; tibia and tarsus ochre. Tissues of tergalii uniformly grey, without pigmentation associated with trachea, so tracheae poorly visible (Figs 63 f, 64 a, b). Head. Antenna (Fig. 62 a). Length ~ 1.5 × head length. As typical for subgenus. Developing turbinate eyes in last instar male larvae. Unknown. Labrum (Fig. 62 a) widened distally; long setae on dorsal surface numerous and forming integral, regular transverse row; each seta consists of stout stem and numerous long processes on both sides; setae and their processes intensively yellowish. Right mandible (Fig. 62 a). As typical for subgenus. Left mandible (Fig. 62 a). As typical for subgenus. Hypopharynx (Fig. 62 c) apically evenly covered with setae-like spines. Maxilla (Fig. 62 b). Maxillary palp as long as galea-lacinia. Otherwise, as typical for genus. Labium (Fig. 62 d). Paraglossa with proximal 2 / 3 parallel-sided; three apical setal rows straight and continued by straight row on ventral side of paraglossa. Glossa as long as 3 / 4 of paraglossa, with slender distal portion twice longer than wide proximal portion; distal portion narrowed proximally, widened at middle, with lateral margin convex. Glossa with several long setae in distal 1 / 2 and with several long seta near middle of ventral side. Labial palp without distomedian projection on segment II; segment III cone-shaped, with median margin as long as lateral margin. Thorax. Sterna. With small protuberances on sides of prosternum and close to openings of mesothoracic and metathoracic sternal apodemes (as in Fig. 108 a). Terga (Fig. 63 d) without long setae on midline. Metanotum with unpaired, moderately long, pointed, spine-like protuberance close to posterior margin; without hind protoptera or their vestiges. Legs (Fig. 64 a – f). Hind leg unknown. Fore femur widened in proximal part. Femur. Outer side of each femur with numerous long, pointed setae situated densely and irregularly, forming stripe of three or four setae width. Apex of femur with short, stout, pointed, spine-like setae. Tibia. Patella-tibial suture reduced, i. e., smoothed out and not crossing inner side of tibia. Tibia-tarsal condylus turned to anterior side. Anterior side of tibia with numerous long setae situated densely and irregularly, forming stripe of three to four setae width; each seta stout and brown in proximal part, hair-like and colorless in distal part. Tarsus. Anterior side of tarsus with stripe of setae similar to tibia, but smaller setae. Posterior side of each tarsus with regular row of short, stout, oval setae (looking pointed in profile) and one much longer, thinner, pointed seta distad of them. Claw with row of six or seven subequal denticles, without posterior seta. Abdomen. Terga (Fig. 63 b – e) without long setae on midline. Each abdominal tergum I – VIII with unpaired, long, pointed, spine-like protuberance close to posterior margin. Abdominal terga I – VI without denticles on posterior margins; posterior margins of abdominal terga VII – X with small, sharply pointed denticles; some pointed denticles on surface of abdominal terga, including median spines. Tergalii (Fig. 63 f) of abdominal segment I absent; tergalii II – VII oval, nearly subequal, tergalii II and VII slightly smaller than others. Each tergalius with costal and anal ribs narrow, smooth, present on proximal 1 / 2 of tergalius only. Paraproct with margins smooth, lacking denticles. Caudalii (Fig. 63 h) without swimming setae or their vestiges. Paracercus small, conic, non-segmented. Pose of subimaginal gonostyli under larval cuticle. Unknown. Subimago. Texture. On all legs of both sexes, each tarsomere covered mostly with blunt microlepides, with pointed microlepides near apex (as in Fig. 70 i). Imago. Unknown. Judging by hypodermal coloration of mature female larva, female imago has following coloration: head, thorax, and abdomen dorsally dark brown; femur of each leg pair dark brown except distal ¼. Egg. Unknown.	en	Kaltenbach, Thomas, Kluge, Nikita J., Gattolliat, Jean-Luc (2025): Redescription of Papuanatula Lugo-Ortiz & McCafferty (Ephemeroptera, Baetidae), with description of a new subgenus and 20 new species. ZooKeys 1227: 159-347, DOI: 10.3897/zookeys.1227.138100
44836D371795592F96323CCBE3EAB69A.taxon	distribution	Distribution. Indonesia: Sulawesi Island (Fig. 147).	en	Kaltenbach, Thomas, Kluge, Nikita J., Gattolliat, Jean-Luc (2025): Redescription of Papuanatula Lugo-Ortiz & McCafferty (Ephemeroptera, Baetidae), with description of a new subgenus and 20 new species. ZooKeys 1227: 159-347, DOI: 10.3897/zookeys.1227.138100
F4D22BC4BCBD5ACA8A6742D9317FD67A.taxon	description	Figs 65, 66, 67, 68, 69, 70, 71	en	Kaltenbach, Thomas, Kluge, Nikita J., Gattolliat, Jean-Luc (2025): Redescription of Papuanatula Lugo-Ortiz & McCafferty (Ephemeroptera, Baetidae), with description of a new subgenus and 20 new species. ZooKeys 1227: 159-347, DOI: 10.3897/zookeys.1227.138100
F4D22BC4BCBD5ACA8A6742D9317FD67A.taxon	etymology	Etymology. The species name obscura refers to the dark color of male imago (Fig. 70 a, b).	en	Kaltenbach, Thomas, Kluge, Nikita J., Gattolliat, Jean-Luc (2025): Redescription of Papuanatula Lugo-Ortiz & McCafferty (Ephemeroptera, Baetidae), with description of a new subgenus and 20 new species. ZooKeys 1227: 159-347, DOI: 10.3897/zookeys.1227.138100
F4D22BC4BCBD5ACA8A6742D9317FD67A.taxon	diagnosis	Diagnosis. Larva. The following combination of characters distinguishes P. obscura sp. nov. from other species of Papuanatula s. str.: body dorsally with irregular row of long, fine, simple setae on midline; abdominal terga without protuberances; femur with clearly outlined wedge-shaped blank on proximal 1 / 2; abdominal terga mostly brownish, terga I and V – VI brighter; posterior margins of abdominal terga II – IX with short, blunt denticles; tracheation of tergalii poorly visible.	en	Kaltenbach, Thomas, Kluge, Nikita J., Gattolliat, Jean-Luc (2025): Redescription of Papuanatula Lugo-Ortiz & McCafferty (Ephemeroptera, Baetidae), with description of a new subgenus and 20 new species. ZooKeys 1227: 159-347, DOI: 10.3897/zookeys.1227.138100
F4D22BC4BCBD5ACA8A6742D9317FD67A.taxon	description	Description. Larva (Figs 65 – 69). Cuticular coloration. Head, pronotum, mesonotum and metanotum brownish, with darker and paler areas; fore protoptera nearly uniformly brownish (Fig. 66 b – d, k). Thoracic pleura brownish, sterna mostly colorless with some areas pale brownish. Cuticle of femur mostly brownish, with clearly outlined wedge-shape blank on proximal 1 / 2; apex of femur bordered with darker brown (Fig. 66 e – g). Tibia and tarsus mostly from colorless to pale brownish, distal end and outer side of tarsus darker brown (Fig. 66 e – g). Abdominal terga mostly brownish, with lateral areas paler; terga I and V – VI more or less paler than others; sterna mostly colorless (Figs 65 a, 66 b). Cerci uniformly pale brownish. Hypodermal coloration. Legs without hypodermal markings. Each abdominal tergum I – IX with wide dark brown transverse band close to anterior margin and with narrower dark brown transverse band close to posterior margin (Fig. 65 a). Tissues of tergalii colorless, without pigmentation associated with trachea, so tracheae poorly visible (Fig. 66 i). Head. Long, fine, soft, colorless setae irregularly situated along midline (Fig. 65 b). Antenna (Fig. 66 c, j). Length ~ 1.5 × head length. As typical for subgenus. Developing turbinate eyes in last instar male larva (Fig. 66 h) with larger facets in middle and smaller facets on periphery; ovoid, rather small, with big distance to each other. Labrum (Fig. 67 a) widened distally; long setae on dorsal surface numerous and forming integral, regular transverse row; each seta consists of stout stem and numerous long processes on both sides; setae and their processes intensively yellowish colored. Right mandible (Fig. 67 d). As typical for subgenus. Left mandible (Fig. 67 c). As typical for subgenus. Hypopharynx (Fig. 67 b). As typical for genus. Maxilla (Fig. 67 b). Maxillary palp as long as galea-lacinia. Otherwise, as typical for the genus. Labium (Fig. 67 e – g). Paraglossae widened near middle, with lateral side forming concavity in proximal part; three apical setal rows sharply bent at apex of paraglossa. Glossa shorter than half of paraglossa, with finger-like (distal) portion as long as triangular (proximal) portion. Glossa with several long setae at apex and one long seta near middle of ventral side. Labial palp without distomedian projection on segment II; segment III with median margin longer than lateral margin. Thorax. Sterna. With small protuberances on sides of prosternum and close to openings of mesothoracic and metathoracic sternal apodemes (as in Fig. 108 a). Terga (Fig. 65 b). Long, fine, soft, colorless setae irregularly situated along midline of all terga. Metanotum without hind protoptera or their vestiges. Legs (Fig. 68 a – f). Fore femur slightly widened in proximal part; hind tibia shorter than others. Femur. Outer side of each femur with single regular row of long, hair-like setae bearing numerous fine, short branches on all sides. Tibia. Patella-tibial suture present on all legs, terminated near middle of inner margin of tibia. Tibia-tarsal condylus turned to anterior side. Anterior side of each tibia with regular row of setae similar to that on femur. Tarsus. Anterior side of each tarsus with regular row of similar, but smaller (shorter and narrower) setae. Posterior side of each tarsus with regular row of short, stout, oval setae (looking pointed in profile) and one much longer, thinner, pointed seta distad of them. Claw with row of six short denticles and one somewhat larger denticle distad of them; one long, arched posterior setae (Fig. 68 e); occasionally 2 such setae (Fig. 68 c). Abdomen. Terga (Figs 65 b, 69 b – d). Long, fine, soft, colorless setae irregularly situated along midline of all abdominal terga. Abdominal terga without dorsal unpaired or paired protuberances, only with slightly expressed paired, submedian elevations. Abdominal terga with small, roundish scales with small sockets and radial striation. Posterior margins of abdominal terga II – IX with short, blunt denticles. Posterior margin of tergum X with smaller, blunt denticles. Tergalii (Fig. 66 i) of abdominal segment I absent; tergalii II – VII subequal, oval. Each tergalius with costal and anal ribs narrow, smooth, present on proximal 1 / 2 of tergalius only. Paraproct (Fig. 69 a) with posterior prolongation bent toward bases of caudalii, with many small, equal denticles on median-posterior margin (Kluge and Novikova 2014: fig. 37). Caudalii (Fig. 66 a) without swimming setae; vestiges of swimming setae present on distal part of cerci. Paracercus short, consisting of ~ 6 – 8 segments. Pose of subimaginal gonostyli under larval cuticle (Fig. 71 a). In mature larva ready to molt to subimago, subimaginal gonostyli packed under larval cuticle in “ Labiobaetis - type ” pose: 2 nd segments directed medially and bent proximally; 3 rd segment directed medially (as continuation of 2 nd segment) and narrowed apically, being deformed corresponding to space between subimaginal styliger and larval cuticle. Subimago. Cuticular coloration. Pronotum and prosternum partly brown (as in Fig. 60 f). Mesonotum pale brown with medioparapsidal suture colorless, other sutures darker brown (Fig. 70 d). Meso- and metathoracic pleura and sterna with colorless, pale brownish and dark brown areas (Fig. 70 e). Cuticle of wings colorless, with microtrichiae brownish. Legs nearly colorless, with pale brown bordering on femur and base of tibia (as in Fig. 36 d). Abdomen diffusely colored with very pale brownish, mostly in distal part. Cerci colorless with setae brown. Hypodermal coloration. As in imago. Texture. (Fig. 70 i). On all legs of both sexes, each tarsomere covered mostly with blunt microlepides, with pointed microlepides near apex. Imago. Imago, male (Fig. 70 a – c, f). Head dark brown. Antennae ochre. Turbinate eyes dark brown, high and narrow, cylindrical, with faceted surfaces round and widely separated. Thorax brown, equally dark dorsally, laterally, and ventrally. Fore wing with membrane mostly colorless, area of pterostigma slightly tinged with brownish; veins intensively ochre-brownish. Pterostigma with 5 – 10 oblique cross veins (Fig. 70 a). Fore leg mostly pale brown, middle and hind legs mostly ochre; femur of each leg with brown apex (Fig. 70 f). Abdominal coloration similar to that of female and larva, but darker: all terga and sterna brown, each tergum I – IX with wide darker brown transverse band close to anterior margin and with narrower dark brown transverse band close to posterior margin (Fig. 70 a – c). Cerci brown. Genitalia (Fig. 71 b, c). Sterno-styligeral muscle absent. Each unistyliger slightly widened apically, with median margin concave and apex thickened. At lateral side of gonostylus, its 1 st segment roundly-convex at apex and separated from 2 nd segment by concavity; at median side of gonostylus, 1 st segment more gradually turns to 2 nd segment. Second segment equally wide along its length. Third (terminal) segment of gonostylus nearly as wide as 2 nd, with length twice exceeding width. Penial bridge with poorly expressed membranous projection between unistyligers. Each gonovectis semicircular, with lateral (basal) and median (apical) portions equally long, apex bent medially. Imago, female (Fig. 70 g, h). Head and thorax dorsally dark brown, ventrally mostly ochre. Hypodermal abdominal coloration as in larva: mostly ochre, each tergum I – IX with wide dark brown transverse band close to anterior margin and with narrower dark brown transverse band close to posterior margin. Coloration of legs, wings, and cerci as in male. Egg (Fig. 37 d, e). Irregularly oval. Chorion without regular relief.	en	Kaltenbach, Thomas, Kluge, Nikita J., Gattolliat, Jean-Luc (2025): Redescription of Papuanatula Lugo-Ortiz & McCafferty (Ephemeroptera, Baetidae), with description of a new subgenus and 20 new species. ZooKeys 1227: 159-347, DOI: 10.3897/zookeys.1227.138100
F4D22BC4BCBD5ACA8A6742D9317FD67A.taxon	distribution	Distribution. New Guinea (Fig. 147).	en	Kaltenbach, Thomas, Kluge, Nikita J., Gattolliat, Jean-Luc (2025): Redescription of Papuanatula Lugo-Ortiz & McCafferty (Ephemeroptera, Baetidae), with description of a new subgenus and 20 new species. ZooKeys 1227: 159-347, DOI: 10.3897/zookeys.1227.138100
A4617B4D05B95EF280888D8E2A29A34B.taxon	description	Figs 72, 73, 74, 75, 76	en	Kaltenbach, Thomas, Kluge, Nikita J., Gattolliat, Jean-Luc (2025): Redescription of Papuanatula Lugo-Ortiz & McCafferty (Ephemeroptera, Baetidae), with description of a new subgenus and 20 new species. ZooKeys 1227: 159-347, DOI: 10.3897/zookeys.1227.138100
A4617B4D05B95EF280888D8E2A29A34B.taxon	etymology	Etymology. The species name obscurella refers to the dark color of male imago (Fig. 75 d).	en	Kaltenbach, Thomas, Kluge, Nikita J., Gattolliat, Jean-Luc (2025): Redescription of Papuanatula Lugo-Ortiz & McCafferty (Ephemeroptera, Baetidae), with description of a new subgenus and 20 new species. ZooKeys 1227: 159-347, DOI: 10.3897/zookeys.1227.138100
A4617B4D05B95EF280888D8E2A29A34B.taxon	diagnosis	Diagnosis. Larva. The following combination of characters distinguishes P. obscurella sp. nov. from other species of Papuanatula s. str.: body without long, fine, simple setae along midline; abdominal terga without protuberances; femur without hypodermal pigmentation; abdominal terga with pointed denticles on posterior margins; tergalii with non-pigmented tracheae; abdomen with hypodermal coloration (brown band on posterior margins of terga I – IX); turbinate eyes with diminished facetted surfaces.	en	Kaltenbach, Thomas, Kluge, Nikita J., Gattolliat, Jean-Luc (2025): Redescription of Papuanatula Lugo-Ortiz & McCafferty (Ephemeroptera, Baetidae), with description of a new subgenus and 20 new species. ZooKeys 1227: 159-347, DOI: 10.3897/zookeys.1227.138100
A4617B4D05B95EF280888D8E2A29A34B.taxon	description	Description. Larva (Figs 72 – 74). Cuticular coloration. Head, pronotum, mesonotum and metanotum brownish, with darker and paler areas; fore protoptera nearly uniformly brownish (Fig. 72 a, e, f). Thoracic pleura brownish, sterna mostly colorless. Cuticle of femur with wedge-shape colorless blank on proximal 1 / 2 and colorless blank occupying most part of distal 1 / 2; other surface of femur brownish, apex bordered with darker brown (Fig. 72 b – d). Tibia and tarsus from colorless to pale brownish (Fig. 72 b – d). Abdominal terga mostly brownish, with lateral areas paler; terga V – VI more or less paler than others; sterna mostly colorless (Fig. 72 a). Cerci uniformly pale brownish. Hypodermal coloration. Legs without hypodermal markings. Hypodermal coloration of abdomen either non-expressed or represented by dark brown transverse band close to posterior margin of each tergum I – IX, sometime with other brown markings on abdominal terga (Fig. 72 h, i). Tissues of tergalii colorless, without pigmentation associated with trachea, so tracheae poorly visible (Fig. 72 g – i). Head. Antenna (Fig. 72 e). Length ~ 2 × head length. As typical for subgenus. Developing turbinate eyes of last instar male larva (Fig. 72 e) with larger facets in middle and smaller facets on periphery. Labrum (Fig. 73 e) widened distally; long setae on dorsal surface numerous and forming integral, regular transverse row; each seta consists of stout stem and numerous long processes on both sides. Right mandible (Fig. 73 b, d). As typical for subgenus. Left mandible (Fig. 73 a, c). As typical for subgenus. Hypopharynx (Fig. 73 f). As typical for genus. Maxilla. (Fig. 73 h). Maxillary palp as long as galea-lacinia. Otherwise, as typical for genus. Labium. (Fig. 73 g) Paraglossae widened near middle, with lateral side forming concavity in proximal part; three apical setal rows sharply bent at apex of paraglossa. Glossa shorter than half of paraglossa, with finger-like (distal) portion as long as triangular (proximal) portion. Glossa with several long setae at apex and one long seta near middle of ventral side. Labial palp without distomedian projection on segment II. Thorax. Sterna. Without protuberances. Terga without protuberances; without long, fine setae on midline. Metanotum without hind protoptera or their vestiges. Legs (Figs 72 b – d, 74 a, b). Fore femur slightly widened in proximal part; hind tibia shorter than others. Femur. Outer side of each femur with single regular row of long, hair-like setae bearing numerous fine, short branches on all sides (as in Figs 41 g, 68 b). Tibia. Patella-tibial suture present on all legs, terminated near middle of inner margin of tibia. Tibia-tarsal condylus turned to anterior side. Anterior side of each tibia with regular row of setae similar to that on femur. Tarsus. Anterior side of each tarsus with regular row of similar, but smaller (shorter and narrower) setae. Posterior side of each tarsus with regular row of short, stout, oval setae (looking pointed in profile) and one much longer, thinner, pointed seta distad of them. Claw with row of four or five short denticles and one somewhat larger denticle distad of them; long, arched posterior seta. Abdomen. Terga (Figs 72 a, 74 c) without dorsal unpaired or paired protuberances, only with slightly expressed, unpaired, median elevations; without long, fine setae on midline. Abdominal terga and sterna without scales. Posterior margins of abdominal terga I – IX with short denticles, short and blunt on several anterior terga, longer and pointed on several posterior terga. Posterior margin of tergum X with very small denticles. Tergalii (Fig. 72 a, g – i) of abdominal segment I absent; tergalii II – VII subequal. Each tergalius with costal and anal ribs narrow, smooth, present on proximal 1 / 2 of tergalius only (as in Fig. 66 i). Paraprocts (Fig. 74 d) without posterior projection; margins membranous, smooth, lacking denticles. Caudalii (Fig. 72 a) without swimming setae or their vestiges. Paracercus short, consisting of ~ 6 – 8 segments. Pose of subimaginal gonostyli under larval cuticle (Fig. 76 b). In mature larva ready to molt to subimago, subimaginal gonostyli packed under larval cuticle in “ Labiobaetis - type ” pose: 2 nd segments directed medially and bent proximally; 3 rd segment directed medially (as continuation of 2 nd segment) and narrowed apically, being deformed corresponding to space between subimaginal styliger and larval cuticle. Subimago. Cuticular coloration. Pronotum and prosternum partly brown (as in Fig. 60 f). Mesonotum pale brown with medioparapsidal suture colorless, other sutures darker brown (Fig. 75 g). Meso- and metathoracic pleura and sterna with colorless, pale brownish and dark brown areas (Fig. 75 h). Cuticle of wings colorless, with microtrichiae brownish. Legs nearly colorless, with pale brown bordering on femur and base of tibia (as in Fig. 36 d). Abdomen diffusely colored with very pale brownish, mostly in distal part. Cerci colorless with setae brown. Hypodermal coloration. As in imago. Texture. On all legs of both sexes, each tarsomere covered mostly with blunt microlepides, with pointed microlepides near apex (as in Fig. 70 i). Imago. Imago, male (Fig. 75 c – f). Head dark brown. Antennae with scape ochre, pedicel brown, flagellum ochre. Turbinate eyes dark brown, high and narrow, cylindrical, with faceted surfaces round and widely separated. Thorax brown, equally dark dorsally, laterally, and ventrally. Fore wing with membrane colorless, veins ochre. Pterostigma with three or four incomplete, oblique cross veins (Fig. 75 f). Legs ochre (Fig. 75 i, j). Abdominal terga brown, each tergum I – IX with darker brown transverse band close to posterior margin; sterna slightly paler, ochre-brown (Fig. 75 c – e). Cerci brown. Genitalia (Fig. 76 a): Sterno-styligeral muscle absent. Each unistyliger parallel-sided, equally wide at base and at apex. At lateral side of gonostylus, its 1 st segment roundly-convex at apex and separated from 2 nd segment by concavity; at median side of gonostylus, 1 st segment gradually turns to 2 nd segment. Second segment equally wide all over its length. 3 rd (terminal) segment of gonostylus nearly as wide as 2 nd, with length slightly exceeding width. Penial bridge with truncated trapezoid projection between unistyligers. Gonovectes dark brown. Each gonovectis parabolic, with lateral (basal) and median (apical) portions equally long, apex slightly bent medially. Imago, female (Fig. 75 a, b). Head and thorax dorsally ochre-brown, ventrally mostly ochre. Abdomen mostly ochre, each tergum I – IX with dark brown transverse band close to posterior margin. Coloration of legs, wings, and cerci as in male. Egg (Fig. 37 c). Irregularly oval. Chorion without regular relief.	en	Kaltenbach, Thomas, Kluge, Nikita J., Gattolliat, Jean-Luc (2025): Redescription of Papuanatula Lugo-Ortiz & McCafferty (Ephemeroptera, Baetidae), with description of a new subgenus and 20 new species. ZooKeys 1227: 159-347, DOI: 10.3897/zookeys.1227.138100
A4617B4D05B95EF280888D8E2A29A34B.taxon	distribution	Distribution. New Guinea (Fig. 147).	en	Kaltenbach, Thomas, Kluge, Nikita J., Gattolliat, Jean-Luc (2025): Redescription of Papuanatula Lugo-Ortiz & McCafferty (Ephemeroptera, Baetidae), with description of a new subgenus and 20 new species. ZooKeys 1227: 159-347, DOI: 10.3897/zookeys.1227.138100
D638F3D2F20A5E3886580FBAF1717051.taxon	description	Figs 77, 78, 79, 80, 81, 82	en	Kaltenbach, Thomas, Kluge, Nikita J., Gattolliat, Jean-Luc (2025): Redescription of Papuanatula Lugo-Ortiz & McCafferty (Ephemeroptera, Baetidae), with description of a new subgenus and 20 new species. ZooKeys 1227: 159-347, DOI: 10.3897/zookeys.1227.138100
D638F3D2F20A5E3886580FBAF1717051.taxon	etymology	Etymology. The species name parabessa refers to the morphological similarity with P. bessa.	en	Kaltenbach, Thomas, Kluge, Nikita J., Gattolliat, Jean-Luc (2025): Redescription of Papuanatula Lugo-Ortiz & McCafferty (Ephemeroptera, Baetidae), with description of a new subgenus and 20 new species. ZooKeys 1227: 159-347, DOI: 10.3897/zookeys.1227.138100
D638F3D2F20A5E3886580FBAF1717051.taxon	diagnosis	Diagnosis. Larva. The following combination of characters distinguishes P. parabessa sp. nov. from other species of Papuanatula s. str.: body dorsally with row of long, fine, simple setae along midline; metanotum and abdominal terga I – V with medioposterior, broad, paired humps; abdominal terga II – VI with paired, semicircular, dark brown markings; femur grey-brown, basally with wedge-shaped blank, dorsally with submarginal, dark grey-brown streak; paracercus with nine segments.	en	Kaltenbach, Thomas, Kluge, Nikita J., Gattolliat, Jean-Luc (2025): Redescription of Papuanatula Lugo-Ortiz & McCafferty (Ephemeroptera, Baetidae), with description of a new subgenus and 20 new species. ZooKeys 1227: 159-347, DOI: 10.3897/zookeys.1227.138100
D638F3D2F20A5E3886580FBAF1717051.taxon	description	Description. Larva (Figs 77 – 82). Body length 4.2 – 5.8 mm, cerci much longer than body length (~ 2 ×). Cuticular coloration (Fig. 77 a – d) Head and thorax dorsally grey-brown, with complex pattern. Abdomen dorsally brown, terga V, VI, and X pale brown; terga II – VI with paired, semicircular, dark brown markings; terga VII – IX with small, paired, dark brown spots. Femur grey-brown, basally with wedge-shaped blank, dorsally with submarginal, dark grey-brown streak. Tibia pale grey-brown; tarsus brown. Head, thorax and abdominal segment I ventrally ecru; protuberances of thoracic sterna brown. Abdominal segments II – VI and X yellow-brown, VII – IX brown. Cerci pale brown. Hypodermal coloration (Fig. 77 a, c). Abdominal terga with narrow, dark brown to blackish, transverse band along posterior margin. Head (Fig. 77 b). Dorsally with irregular row of long, fine, simple setae along midline. Antenna (Fig. 80 h). Length ~ 1.5 × head length. As typical for genus. Developing turbinate eyes in last instar male larva (Fig. 80 h) large, subquadrangular, broadly touching each other. Labrum (Fig. 78 a, b). Length 0.5 × maximum width, laterally convex. Dorsal, sub-marginal arc with ~ 32 feathered setae. Right mandible (Fig. 78 d, e). Margin between prostheca and mola with few denticles. Otherwise, as typical for subgenus. Left mandible (Fig. 78 f, g). Margin between prostheca and mola smooth, without denticles. Otherwise, as typical for subgenus. Hypopharynx (Fig. 78 c). As typical for genus. Maxilla (Fig. 79 c, d). Maxillary palp subequal in length to galea-lacinia; palp segment II approximately as long as segment I. Otherwise, as typical for genus. Labium (Fig. 79 a, b). As typical for the genus. Paraglossa dorsally with two spine-like setae near inner, distolateral margin. Labial palp with segment I 0.9 × length of segments II and III combined. Segment II with minute distomedial protuberance, dorsally with row of five spine-like setae near outer, distolateral margin. Segment III conical, pointed; 0.7 × length of segment II; inner dorsal margin with few feathered setae. Thorax. Sterna. With small protuberances on sides of prosternum and close to openings of mesothoracic and metathoracic sternal apodemes (as in Fig. 108 a). Terga (Figs 77 b, 81 a) with irregular row of long, fine, simple setae along midline. Metanotum with medioposterior, broad, paired humps. Legs (Fig. 88 a – g). Ratio of leg segments: fore leg 0.8: 1.0: 0.3: 0.1, middle leg 0.9: 1.0: 0.3: 0.1 and hind leg 1.0: 1.0: 0.3: 0.2. Femur. Length ~ 4 × maximum width. Claw with one row of seven or eight denticles and one posterior seta. Abdomen. Terga (Figs 77 b, 81 a, b, 82 d, e) with irregular row of long, fine, simple setae along midline. Terga I – V with medioposterior, broad, paired humps. Posterior margin of terga: I – II smooth, without denticles; III – IX with triangular, apically rounded denticles, partly with minute, pointed denticles in between. Surface with scattered small, ovoid, striated scales. Tergalii (Fig. 81 e, f). Ovoid, tracheation well developed; margins smooth, with short, fine, simple setae. Paraproct (Fig. 82 a – c). Posterior margin with prolongation and row of minute denticles. Caudalii (Fig. 81 c, d). Cerci apart from basal and distal part with 1 – 5 swimming setae per segment, initially increasing and then again decreasing toward distal part. Paracercus with nine segments. Pose of subimaginal gonostyli under larval cuticle (Fig. 80 i). In mature larva ready to molt to subimago, subimaginal gonostyli packed under larval cuticle in “ Labiobaetis - type ” pose: 2 nd segments directed medially and bent proximally; 3 rd segment directed medially (as continuation of 2 nd segment) and narrowed apically. Subimago. Unknown. Imago. Unknown. Egg. Unknown.	en	Kaltenbach, Thomas, Kluge, Nikita J., Gattolliat, Jean-Luc (2025): Redescription of Papuanatula Lugo-Ortiz & McCafferty (Ephemeroptera, Baetidae), with description of a new subgenus and 20 new species. ZooKeys 1227: 159-347, DOI: 10.3897/zookeys.1227.138100
D638F3D2F20A5E3886580FBAF1717051.taxon	distribution	Distribution. New Guinea (Fig. 147).	en	Kaltenbach, Thomas, Kluge, Nikita J., Gattolliat, Jean-Luc (2025): Redescription of Papuanatula Lugo-Ortiz & McCafferty (Ephemeroptera, Baetidae), with description of a new subgenus and 20 new species. ZooKeys 1227: 159-347, DOI: 10.3897/zookeys.1227.138100
5D94675EDBEE563F91DF3A37F5829D8D.taxon	description	Figs 83, 84, 85, 86, 87, 88, 89	en	Kaltenbach, Thomas, Kluge, Nikita J., Gattolliat, Jean-Luc (2025): Redescription of Papuanatula Lugo-Ortiz & McCafferty (Ephemeroptera, Baetidae), with description of a new subgenus and 20 new species. ZooKeys 1227: 159-347, DOI: 10.3897/zookeys.1227.138100
5D94675EDBEE563F91DF3A37F5829D8D.taxon	etymology	Etymology. The species name paracopis refers to the morphological similarity with P. copis.	en	Kaltenbach, Thomas, Kluge, Nikita J., Gattolliat, Jean-Luc (2025): Redescription of Papuanatula Lugo-Ortiz & McCafferty (Ephemeroptera, Baetidae), with description of a new subgenus and 20 new species. ZooKeys 1227: 159-347, DOI: 10.3897/zookeys.1227.138100
5D94675EDBEE563F91DF3A37F5829D8D.taxon	diagnosis	Diagnosis. Larva. The following combination of characters distinguishes P. paracopis sp. nov. from other species of Papuanatula s. str.: body dorsally without row of long, fine, simple setae along midline; metanotum and abdominal terga I – VIII medially with conspicuous, long, hook-like protuberance, bent posteriad, on abdominal segment (s) IX (X) vestigial; labial palp segment II with small, distolateral protuberance; segment III oblong, conical; femur anteriorly with angulate blank in basal 1 / 2; paracercus vestigial.	en	Kaltenbach, Thomas, Kluge, Nikita J., Gattolliat, Jean-Luc (2025): Redescription of Papuanatula Lugo-Ortiz & McCafferty (Ephemeroptera, Baetidae), with description of a new subgenus and 20 new species. ZooKeys 1227: 159-347, DOI: 10.3897/zookeys.1227.138100
5D94675EDBEE563F91DF3A37F5829D8D.taxon	description	Description. Larva (Figs 83 – 89). Body length ~ 6.8 mm, cerci much longer than body length. Cuticular coloration (Figs 83 a – c, 84 a, b). Head, thorax and abdomen dorsally brown to dark brown, abdominal segments X and partly IX brighter. Thorax with complex markings. Abdominal segments I – IX anteriorly and laterally darker, segments IV – IX with two dark brown dots in anteromedial area. Head and thorax ventrally ecru, thorax with pale brown to dark brown, paired protuberances near distolateral corners of sterna; abdomen ventrally pale yellow-brown, laterally with brown markings. Legs yellow-brown to brown, femur medially with submarginal brown streak and basally with angulate blank. Caudalii yellow-brown. Hypodermal coloration (Figs 83 a, 84 a). Abdominal terga I – IX with wide dark brown transverse band close to anterior margin and narrower dark brown transverse band close to posterior margin. Head. Antenna (Fig. 86 d). Length ~ 1.5 × head length. As typical for subgenus. Developing turbinate eyes in last instar male larva (Fig. 86 d) rather narrow, with big distance to each other. Labrum (Fig. 85 a, b). Length 0.6 × maximum width, laterally convex. Dorsal, sub-marginal arc with> 50 densely articulated, feathered setae. Right mandible (Fig. 85 d, e). Margin between prostheca and mola smooth, without denticles. Otherwise, as typical for subgenus. Left mandible (Fig. 85 f – h). Margin between prostheca and mola with minute denticles toward subtriangular process. Otherwise, as typical for subgenus. Hypopharynx (Fig. 85 c). As typical for genus. Maxilla (Fig. 86 c). Maxillary palp slightly longer than galea-lacinia; palp segment II slender, partly sclerotized, approx. 1.2 × as long as segment I, segment I thicker than segment II. Otherwise, as typical for genus. Labium (Fig. 86 a, b). As typical for genus. Paraglossa dorsally with two spine-like setae near inner, distolateral margin. Labial palp with segment I 0.8 × length of segments II and III combined. Segment II with very small, rounded, distomedial protuberance, dorsally with row of four or five spine-like setae near outer, distolateral margin. Segment III oblong, pointed, 0.6 × length of segment II; inner dorsal margin with few feathered setae. Thorax. Sterna. Protuberances poorly developed. Terga (Figs 83 b, 84 b, 88 a, 89 a). Mesonotum with small, posteromedial protuberance; metanotum posteromedially with hook-like, pointed protuberance, bent posteriorly. Immature larva with short, acute, posteromedial protuberance on pronotum. Surface of fore protoptera along developing veins and on inner margins with small, triangular, pointed scales. Legs (Fig. 87 a – e). Ratio of leg segments: fore leg 0.9: 1.0: 0.3: 0.2, middle leg 1.0: 1.0: 0.3: 0.2 and hind leg 1.1: 1.0: 0.3: 0.2. Femur. Length ~ 3.5 × maximum width. Claw with one row of nine denticles, 1 st denticle longer than other ones, and one posterior seta. Abdomen. Terga (Figs 83 b, 84 b, 88 a – c, 89 a – e). Abdominal terga I – VIII posteromedially with conspicuous, long, hook-like protuberance, bent posteriad, on abdominal segment (s) IX (X) vestigial; Posterior margin of terga: I smooth, without spines; II – IX with small, triangular, pointed denticles, increasing in length toward IX. Surface with scattered small, triangular, pointed scales. Tergalii (Fig. 87 f, g). Broad oblique ovoid; tracheation developed, hardly visible due to brown pigmentation in large middle part; margins with minute serration and many short, fine, simple setae. Paraproct (Fig. 88 g). Posterior margin with prolongation and with row of many minute denticles. Caudalii (Fig. 88 d – f). Cerci without swimming setae; sometimes few rudimentary swimming setae or insertions still present. Paracercus vestigial. Pose of subimaginal gonostyli under larval cuticle. Unknown. Subimago. Unknown. Imago. Unknown. Egg. Unknown.	en	Kaltenbach, Thomas, Kluge, Nikita J., Gattolliat, Jean-Luc (2025): Redescription of Papuanatula Lugo-Ortiz & McCafferty (Ephemeroptera, Baetidae), with description of a new subgenus and 20 new species. ZooKeys 1227: 159-347, DOI: 10.3897/zookeys.1227.138100
5D94675EDBEE563F91DF3A37F5829D8D.taxon	distribution	Distribution. New Guinea (Fig. 147).	en	Kaltenbach, Thomas, Kluge, Nikita J., Gattolliat, Jean-Luc (2025): Redescription of Papuanatula Lugo-Ortiz & McCafferty (Ephemeroptera, Baetidae), with description of a new subgenus and 20 new species. ZooKeys 1227: 159-347, DOI: 10.3897/zookeys.1227.138100
7506516145BE5F8DA017EF837EBC172B.taxon	description	Figs 90, 91, 92, 93, 94, 95, 96	en	Kaltenbach, Thomas, Kluge, Nikita J., Gattolliat, Jean-Luc (2025): Redescription of Papuanatula Lugo-Ortiz & McCafferty (Ephemeroptera, Baetidae), with description of a new subgenus and 20 new species. ZooKeys 1227: 159-347, DOI: 10.3897/zookeys.1227.138100
7506516145BE5F8DA017EF837EBC172B.taxon	etymology	Etymology. The species name paralenos refers to the morphological similarity with P. lenos.	en	Kaltenbach, Thomas, Kluge, Nikita J., Gattolliat, Jean-Luc (2025): Redescription of Papuanatula Lugo-Ortiz & McCafferty (Ephemeroptera, Baetidae), with description of a new subgenus and 20 new species. ZooKeys 1227: 159-347, DOI: 10.3897/zookeys.1227.138100
7506516145BE5F8DA017EF837EBC172B.taxon	diagnosis	Diagnosis. Larva. The following combination of characters distinguishes P. lenos from other species of Papuanatula s. str.: body dorsally without row of long, fine, simple setae along midline; body dorsally without protuberances; thorax ventrally without protuberances; pronotum dorsally with large, medial, dark brown marking at anterior margin, and fine, dark brown band along posterior margin; mesothorax with dark brown band along anterior margin; femur anteriorly with large, oblong, red brown to dark brown marking in basal 1 / 2, and pale red brown to dark brown marking in mediodistal area; paracercus with 6 – 8 segments; paraproct without posterior prolongation.	en	Kaltenbach, Thomas, Kluge, Nikita J., Gattolliat, Jean-Luc (2025): Redescription of Papuanatula Lugo-Ortiz & McCafferty (Ephemeroptera, Baetidae), with description of a new subgenus and 20 new species. ZooKeys 1227: 159-347, DOI: 10.3897/zookeys.1227.138100
7506516145BE5F8DA017EF837EBC172B.taxon	description	Description. Larva (Figs 90 – 96). Body length 4.0 – 5.5 mm, cerci much longer than body length. Cuticular coloration (Figs 90 a – c, 91 a). Head and thorax dorsally yellow-brown to grey-brown; Abdomen dorsally yellow-brown to dark brown; segments II – IV entirely dark brown, or segment IV with dark brown, crown-like marking; segments I – IX with oblique, dark brown to blackish lateral markings. Head and thorax ventrally ecru, abdomen ventrally pale yellow-brown. Legs yellow-brown to brown; Caudalii yellow-brown. Hypodermal coloration (Figs 90 a – c, 91 a). Pronotum with large, dark brown marking medially at anterior margin and fine, dark brown band along posterior margin; mesothorax with narrow, dark brown band along anterior margin. Abdominal terga I – IX with wide dark brown transverse band close to anterior margin and narrower dark brown transverse band close to posterior margin. Femur anteriorly with large, oblong, red brown to dark brown marking in basal 1 / 2 (somewhat variable in shape), and pale red brown to dark brown marking in mediodistal area; posteriorly with two red-brown to dark brown streaks close to dorsal margin. Head. Antenna (Fig. 90 a, c). Length ~ 2 × head length. As typical for subgenus. Developing turbinate eyes in last instar male larva (Fig. 93 e) large, subquadrangular, nearly touching each other in the middle. Labrum (Figs 92 a, b, 96 f). Length 0.5 × maximum width, laterally convex. Dorsal, sub-marginal arc with 15 – 17 feathered setae. Right mandible (Fig. 92 d, e). Margin between prostheca and mola smooth. Otherwise, as typical for subgenus. Left mandible (Fig. 92 f, g). Margin between prostheca and mola smooth, with several spines close to subtriangular process. Otherwise, as typical for subgenus. Hypopharynx (Fig. 92 c). As typical for genus. Maxilla (Fig. 93 c, d). Maxillary palp slightly longer than galea-lacinia; palp segment II slightly shorter than segment I. Otherwise, as typical for genus. Labium (Fig. 93 a, b). As typical for genus. Paraglossa with one spine-like seta on inner, distolateral margin. Labial palp with segment I subequal in length to segments II and III combined. Segment II without distomedial protuberance, dorsally with row of four spine-like setae near outer, distolateral margin. Segment III slightly bulbous, pointed, 0.8 × length of segment II; inner dorsal margin with few feathered setae. Thorax. Sterna without protuberances. Terga without protuberances. Legs (Fig. 94 a – h). Ratio of leg segments: fore leg 0.9: 1.0: 0.3: 0.1, middle leg 1.0: 1.0: 0.3: 0.1 and hind leg 1.1: 1.0: 0.3: 0.1. Femur. Length ~ 2.6 × maximum width. Many medium, pointed, spine-like setae along ventral margin. Claw with one row of 5 – 7 denticles, distalmost denticle with distance to other denticles; one posterior seta. Abdomen. Terga (Figs 95 a – c, 96 a – e). Abdominal terga without protuberances. Posterior margin of terga: I – III smooth, without denticles; IV – IX with triangular, pointed denticles. Surface with scattered small, trapezoid, striated, apically serrate scales. Tergalii (Fig. 93 f, g). Ovoid, tracheation rather poorly developed; costal margin with minute serration and short, fine, simple setae; anal margin smooth, with short, fine, simple setae. Paraproct (Fig. 95 f). Posterior margin without prolongation, smooth, without denticles. Caudalii (Fig. 95 d, e). Cerci without swimming setae; sometimes one or two insertions still present. Paracercus with 6 – 8 segments. Pose of subimaginal gonostyli under larval cuticle (Fig. 94 i). As typical for subgenus. Segment III conical. Subimago. Unknown. Imago. Unknown. Egg. Unknown.	en	Kaltenbach, Thomas, Kluge, Nikita J., Gattolliat, Jean-Luc (2025): Redescription of Papuanatula Lugo-Ortiz & McCafferty (Ephemeroptera, Baetidae), with description of a new subgenus and 20 new species. ZooKeys 1227: 159-347, DOI: 10.3897/zookeys.1227.138100
7506516145BE5F8DA017EF837EBC172B.taxon	distribution	Distribution. New Guinea (Fig. 147).	en	Kaltenbach, Thomas, Kluge, Nikita J., Gattolliat, Jean-Luc (2025): Redescription of Papuanatula Lugo-Ortiz & McCafferty (Ephemeroptera, Baetidae), with description of a new subgenus and 20 new species. ZooKeys 1227: 159-347, DOI: 10.3897/zookeys.1227.138100
69934095D95F52A181A8F295840862F3.taxon	description	Figs 97, 98, 99, 100, 101, 102	en	Kaltenbach, Thomas, Kluge, Nikita J., Gattolliat, Jean-Luc (2025): Redescription of Papuanatula Lugo-Ortiz & McCafferty (Ephemeroptera, Baetidae), with description of a new subgenus and 20 new species. ZooKeys 1227: 159-347, DOI: 10.3897/zookeys.1227.138100
69934095D95F52A181A8F295840862F3.taxon	etymology	Etymology. The species name paratuber refers to the morphological similarity with P. tuber.	en	Kaltenbach, Thomas, Kluge, Nikita J., Gattolliat, Jean-Luc (2025): Redescription of Papuanatula Lugo-Ortiz & McCafferty (Ephemeroptera, Baetidae), with description of a new subgenus and 20 new species. ZooKeys 1227: 159-347, DOI: 10.3897/zookeys.1227.138100
69934095D95F52A181A8F295840862F3.taxon	diagnosis	Diagnosis. Larva. The following combination of characters distinguishes P. paratuber sp. nov. from other species of Papuanatula s. str.: body dorsally without row of long, fine, simple setae along midline; abdominal terga I – VIII (IX) with medium, pointed, dorsoposteriorly oriented, medial protuberances; pronotum without protuberances; femur with medial, grey marking; paracercus vestigial; body size 4.3 – 5.2 mm.	en	Kaltenbach, Thomas, Kluge, Nikita J., Gattolliat, Jean-Luc (2025): Redescription of Papuanatula Lugo-Ortiz & McCafferty (Ephemeroptera, Baetidae), with description of a new subgenus and 20 new species. ZooKeys 1227: 159-347, DOI: 10.3897/zookeys.1227.138100
69934095D95F52A181A8F295840862F3.taxon	description	Description. Larva (Figs 97 – 102). Body length 4.3 – 5.2 mm, cerci much longer than body length (~ 1.3 ×). Cuticular coloration (Fig. 97 a – d). Head, thorax and abdomen dorsally brown or grey-brown; thorax with complex pattern; abdominal segments II – IV and VII – IX darker, V – VI and X brighter. Legs grey to yellow-brown; femur medially with grey marking, yellow-brown in distal area, blank area in basal part. Head and thorax ventrally pale grey, protuberances on thoracic sterna darker; abdomen yellow-brown to grey, sterna VII and VIII darker. Hypodermal coloration (Fig. 97 a, b). Abdominal segments I – IX dorsally with dark brown, narrow, transverse band on posterior margins; intersegmental membranes slightly reddish-grey. Head. Antenna (Fig. 100 h). Length 1.5 × head length. As typical for subgenus. Developing turbinate eyes in last instar male larva (Fig. 100 h) large, sub quadrangular, nearly touching each other in the middle. Labrum (Fig. 98 a, b). Length 0.5 × maximum width, laterally convex. Dorsal, sub-marginal arc with ~ 9 feathered setae. Right mandible (Fig. 98 d, e). Margin between prostheca and mola smooth. Otherwise, as typical for subgenus. Left mandible (Fig. 98 f, g). Margin between prostheca and mola smooth. Otherwise, as typical for subgenus. Hypopharynx (Fig. 98 c). As typical for genus. Maxilla (Fig. 99 c, d). Maxillary palp ~ 1.2 × length of galea-lacinia; palp segment II ~ 1.1 × length of segment I. Otherwise, as typical for genus. Labium (Fig. 99 a, b). As typical for genus. Paraglossa dorsally with two spine-like setae near inner, distolateral margin. Labial palp with segment I ~ 1.2 × length of segments II and III combined. Segment II with slight, broadly rounded, distomedial protuberance, dorsally with row of four spine-like setae near outer, distolateral margin. Segment III slightly pentagonal, pointed, 0.8 × length of segment II. Thorax. Sterna. With small protuberances on sides of prosternum and close to openings of mesothoracic and metathoracic sternal apodemes (as Fig. 108 a). Terga (Fig. 102 a, d). Pro-, meso- and metanotum without protuberances. Legs (Fig. 100 a – e). Ratio of leg segments: fore leg 0.9: 1.0: 0.4: 0.1, middle leg 0.9: 1.0: 0.3: 0.1 and hind leg 1.0: 1.0: 0.4: 0.2. Femur. Length ~ 3.4 × maximum width; inner margin with short, spine-like setae along margin. Claw with one row of five or six denticles and one posterior seta. Abdomen. Terga (Figs 101 a, b, 102 a – c, e). Terga I – VIII with medium, pointed, dorsoposteriorly oriented, medial protuberances. Posterior margin of terga: I smooth, without denticles; II with rudimentary denticles; III – IX with small, triangular denticles, apically sometimes split. Tergalii (Fig. 101 c, d). Ovoid, tracheation mainly limited to trunk; margins smooth, with short, fine, simple setae. Paraproct (Fig. 101 e). Posterior margin with prolongation and row of minute denticles. Caudalii (Fig. 100 f, g). Cerci without swimming setae. Paracercus vestigial. Pose of subimaginal gonostyli under larval cuticle. Unknown. Subimago. Unknown. Imago. Unknown. Egg. Unknown.	en	Kaltenbach, Thomas, Kluge, Nikita J., Gattolliat, Jean-Luc (2025): Redescription of Papuanatula Lugo-Ortiz & McCafferty (Ephemeroptera, Baetidae), with description of a new subgenus and 20 new species. ZooKeys 1227: 159-347, DOI: 10.3897/zookeys.1227.138100
69934095D95F52A181A8F295840862F3.taxon	distribution	Distribution. New Guinea (Fig. 147).	en	Kaltenbach, Thomas, Kluge, Nikita J., Gattolliat, Jean-Luc (2025): Redescription of Papuanatula Lugo-Ortiz & McCafferty (Ephemeroptera, Baetidae), with description of a new subgenus and 20 new species. ZooKeys 1227: 159-347, DOI: 10.3897/zookeys.1227.138100
43ABA01CE8F55DD4A468D4334657B92B.taxon	description	Figs 103, 104, 105, 106, 107, 108	en	Kaltenbach, Thomas, Kluge, Nikita J., Gattolliat, Jean-Luc (2025): Redescription of Papuanatula Lugo-Ortiz & McCafferty (Ephemeroptera, Baetidae), with description of a new subgenus and 20 new species. ZooKeys 1227: 159-347, DOI: 10.3897/zookeys.1227.138100
43ABA01CE8F55DD4A468D4334657B92B.taxon	etymology	Etymology. The species name parvatubera is based on the Latin words parva tubera meaning “ small humps ”, referring to the specific abdominal protuberances.	en	Kaltenbach, Thomas, Kluge, Nikita J., Gattolliat, Jean-Luc (2025): Redescription of Papuanatula Lugo-Ortiz & McCafferty (Ephemeroptera, Baetidae), with description of a new subgenus and 20 new species. ZooKeys 1227: 159-347, DOI: 10.3897/zookeys.1227.138100
43ABA01CE8F55DD4A468D4334657B92B.taxon	diagnosis	Diagnosis. Larva. The following combination of characters distinguishes P. parvatubera sp. nov. from other species of Papuanatula s. str.: body dorsally without row of long, fine, simple setae along midline; abdominal terga (III) IV – VIII medially with small, triangular protuberance, oriented posteriorly, terga II, III and IX sometimes with vestigial protuberances; femur medially with broad, transversal marking and large blanks in basal and distal area; paracercus with seven or eight segments.	en	Kaltenbach, Thomas, Kluge, Nikita J., Gattolliat, Jean-Luc (2025): Redescription of Papuanatula Lugo-Ortiz & McCafferty (Ephemeroptera, Baetidae), with description of a new subgenus and 20 new species. ZooKeys 1227: 159-347, DOI: 10.3897/zookeys.1227.138100
43ABA01CE8F55DD4A468D4334657B92B.taxon	description	Description. Larva (Figs 103 – 108). Body length 2.8 – 3.1 mm, cerci somewhat shorter than body length (~ 0.8 ×). Cuticular coloration (Fig. 103 a – c). Head, thorax and abdomen dorsally brown to grey-brown; thorax with complex pattern; abdominal segments IV and V laterally pale grey-brown, X pale grey-brown. Femur medially with grey-brown transversal marking, basally and distally with large blanks; tibia ecru with grey-brown in medial area; tarsus distally yellow-brown, basally brown. Head, thorax and abdomen ventrally pale brown to brown, protuberances of thoracic sterna dark brown, abdominal segments IX and X beige. Cerci pale grey-brown. Hypodermal coloration. None (apart from antennal flagellum, as typical for subgenus). Head. Antenna (Fig. 103 a, c). Length 1.5 × head length. As typical for subgenus. Developing turbinate eyes in last instar male larva (Fig. 106 c) large, nearly touching each other, slightly trapezoid. Labrum (Fig. 104 a, b). Relatively small, length 0.5 × maximum width, laterally convex. Dorsal, sub-marginal arc with ~ 10 feathered setae. Right mandible (Fig. 104 d, e). Margin between prostheca and mola with row of minute denticles. Otherwise, as typical for subgenus. Left mandible (Fig. 104 f, g). Margin between prostheca and mola with row of minute denticles. Otherwise, as typical for subgenus. Hypopharynx (Fig. 104 c). As typical for genus. Maxilla (Fig. 105 c, d). Maxillary palp subequal in length to galea-lacinia, robust; palp segment II ~ 1.3 × length of segment I. Otherwise, as typical for genus. Labium (Fig. 105 a, b). As typical for genus. Paraglossa dorsally with two spine-like setae near inner, distolateral margin. Labial palp with segment I ~ 1.1 × length of segments II and III combined. Segment II without distomedial protuberance, dorsally with row of four spine-like setae near outer, distolateral margin. Segment III conical, pointed, 0.7 × length of segment II. Thorax. Sterna (Figs 106 d, 108 a). With small protuberances on sides of prosternum and close to openings of mesothoracic and metathoracic sternal apodemes. Terga without protuberances. Legs (Figs 106 a, b, 108 b). Ratio of leg segments: fore leg 0.9: 1.0: 0.3: 0.2, middle leg 1.0: 1.0: 0.3: 0.2 and hind leg 0.9: 1.0: 0.3: 0.2. Femur. Length ~ 3 × maximum width. Claw with one row of seven denticles, distalmost denticle with distance to other denticles; one posterior seta. Abdomen. Terga (Figs 107 a, b, 108 c – f). Terga (III) IV – VIII posteromedially with small, triangular protuberance, oriented posteriorly, terga II, III and IX sometimes with vestigial protuberance. Posterior margin of terga: I – II smooth, without spines; III – IX with short, rounded denticles, apically carrying minute, fine, acute spines. Surface with scattered small, long-triangular, pointed, striated scales. Tergalii (Fig. 107 d). Narrow oblong, tracheation undeveloped; margins smooth, with few short, fine, simple setae. Paraproct (Fig. 107 c). Posterior margin with prolongation and row of minute denticles. Caudalii (Fig. 106 e – g). Cerci in distal 1 / 2 with 1 – 4 swimming setae per segment. Paracercus with seven or eight segments. Pose of subimaginal gonostyli under larval cuticle (Fig. 107 e). As typical for the subgenus. Subimago. Unknown. Imago. Unknown. Egg. Unknown.	en	Kaltenbach, Thomas, Kluge, Nikita J., Gattolliat, Jean-Luc (2025): Redescription of Papuanatula Lugo-Ortiz & McCafferty (Ephemeroptera, Baetidae), with description of a new subgenus and 20 new species. ZooKeys 1227: 159-347, DOI: 10.3897/zookeys.1227.138100
43ABA01CE8F55DD4A468D4334657B92B.taxon	distribution	Distribution. New Guinea (Fig. 147).	en	Kaltenbach, Thomas, Kluge, Nikita J., Gattolliat, Jean-Luc (2025): Redescription of Papuanatula Lugo-Ortiz & McCafferty (Ephemeroptera, Baetidae), with description of a new subgenus and 20 new species. ZooKeys 1227: 159-347, DOI: 10.3897/zookeys.1227.138100
BBCFDF5234BD518EA41075D89BEB1557.taxon	description	Figs 109, 110, 111, 112, 113	en	Kaltenbach, Thomas, Kluge, Nikita J., Gattolliat, Jean-Luc (2025): Redescription of Papuanatula Lugo-Ortiz & McCafferty (Ephemeroptera, Baetidae), with description of a new subgenus and 20 new species. ZooKeys 1227: 159-347, DOI: 10.3897/zookeys.1227.138100
BBCFDF5234BD518EA41075D89BEB1557.taxon	etymology	Etymology. The species name is based on the Latin word pilosus meaning “ hairy ” and refers to the rows of fine setae on inner margin of femur, outer margin of tibia, and laterally on cerci.	en	Kaltenbach, Thomas, Kluge, Nikita J., Gattolliat, Jean-Luc (2025): Redescription of Papuanatula Lugo-Ortiz & McCafferty (Ephemeroptera, Baetidae), with description of a new subgenus and 20 new species. ZooKeys 1227: 159-347, DOI: 10.3897/zookeys.1227.138100
BBCFDF5234BD518EA41075D89BEB1557.taxon	diagnosis	Diagnosis. Larva. The following combination of characters distinguishes P. pilosa sp. nov. from other species of Papuanatula s. str.: body dorsally without row of long, fine, simple setae along midline; abdominal terga I – VIII (IX) with medium, pointed, dorsoposteriorly oriented, medial protuberances; pronotum with paired, medioposterior protuberances; femur with medial, grey marking; inner margin of femur and outer margin of tibia with irregular rows of medium, fine setae; cerci bilaterally with row of short, fine setae; paracercus vestigial; body size 4.5 mm.	en	Kaltenbach, Thomas, Kluge, Nikita J., Gattolliat, Jean-Luc (2025): Redescription of Papuanatula Lugo-Ortiz & McCafferty (Ephemeroptera, Baetidae), with description of a new subgenus and 20 new species. ZooKeys 1227: 159-347, DOI: 10.3897/zookeys.1227.138100
BBCFDF5234BD518EA41075D89BEB1557.taxon	description	Description. Larva (Figs 109 – 113). Body length 4.5 mm, cerci much longer than body length (~ 1.2 ×). Cuticular coloration (Fig. 109 a – c). Head, thorax and abdomen dorsally reddish-brown; thorax with complex pattern; abdominal segments I, V – VI and X slightly brighter. Legs reddish-brown; femur medially with darker marking, red-brown in distal area, bright area in basal part. Head and thorax ventrally ecru, protuberances on thoracic sterna darker; abdomen yellow-brown. Hypodermal coloration (Fig. 109 a, b). Abdominal segments I – IX dorsally with narrow, dark brown, transverse band on posterior margins; intersegmental membranes slightly reddish-grey. Head. Antenna (Fig. 109 a – c). Length ~ 1.5 × head length. As typical for subgenus. Developing turbinate eyes in last instar male larva unknown. Labrum (Fig. 110 a, b). Length 0.5 × maximum width, laterally convex. Dorsal, sub-marginal arc with ~ 9 feathered setae. Right mandible (Fig. 110 d, e). Margin between prostheca and mola with some minute denticles toward prostheca. Otherwise, as typical for subgenus. Left mandible (Fig. 110 f, g). Margin between prostheca and mola with some minute denticles toward prostheca. Otherwise, as typical for subgenus. Hypopharynx (Fig. 110 c). As typical for genus. Maxilla (Fig. 111 c, d). Maxillary palp slightly longer than galea-lacinia, slender; palp segment II ~ 1.3 × length of segment I. Otherwise, as typical for genus. Labium (Fig. 111 a, b). As typical for the genus. Paraglossa dorsally with two spine-like setae near inner, distolateral margin. Labial palp with segment I ~ 1.2 × length of segments II and III combined. Segment II with slight, broadly rounded, distomedial protuberance, dorsally with row of five spine-like setae near outer, distolateral margin. Segment III slightly pentagonal, pointed, 0.6 × length of segment II. Thorax. Sterna. With small protuberances on sides of prosternum and close to openings of mesothoracic and metathoracic sternal apodemes (as Fig. 108 a). Terga (Figs 109 b, 113 b). Pronotum with paired, blunt, posteromedial protuberances. Metanotum with medium, pointed, dorsally oriented, medial protuberance. Legs (Fig. 112 a – e). Ratio of leg segments: fore leg 1.0: 1.0: 0.3: 0.1, middle leg 1.1: 1.0: 0.3: 0.1 and hind leg 1.2: 1.0: 0.3: 0.1. Femur. Length ~ 3 × maximum width; inner margin with irregular, dense row of medium, fine setae. Tibia. Outer margin with irregular, dense row of medium, fine setae. Claw with one row of eight or nine denticles and one posterior seta. Abdomen. Terga (Fig. 113 a, c – f). Terga I – IX with medium, pointed, dorsoposteriorly oriented, medial protuberances. Posterior margin of terga III – IX with variable, triangular, pointed denticles, spaced on terga III – VI. Tergalii (Fig. 111 f, g). Skew ovoid, tracheation well developed; margins smooth, with short, fine, simple setae. Paraproct (Fig. 111 e). Posterior margin membranous, with prolongation and row of minute denticles. Caudalii (Fig. 112 f – h). Cerci without swimming setae, with bilateral rows of minute setae. Paracercus vestigial. Pose of subimaginal gonostyli under larval cuticle. Unknown. Subimago. Unknown. Imago. Unknown. Egg. Unknown.	en	Kaltenbach, Thomas, Kluge, Nikita J., Gattolliat, Jean-Luc (2025): Redescription of Papuanatula Lugo-Ortiz & McCafferty (Ephemeroptera, Baetidae), with description of a new subgenus and 20 new species. ZooKeys 1227: 159-347, DOI: 10.3897/zookeys.1227.138100
BBCFDF5234BD518EA41075D89BEB1557.taxon	distribution	Distribution. New Guinea (Fig. 147).	en	Kaltenbach, Thomas, Kluge, Nikita J., Gattolliat, Jean-Luc (2025): Redescription of Papuanatula Lugo-Ortiz & McCafferty (Ephemeroptera, Baetidae), with description of a new subgenus and 20 new species. ZooKeys 1227: 159-347, DOI: 10.3897/zookeys.1227.138100
EF608808121A551A880CA064B7D9A843.taxon	description	Figs 17, 18, 19, 20, 21, 22, 23, 24	en	Kaltenbach, Thomas, Kluge, Nikita J., Gattolliat, Jean-Luc (2025): Redescription of Papuanatula Lugo-Ortiz & McCafferty (Ephemeroptera, Baetidae), with description of a new subgenus and 20 new species. ZooKeys 1227: 159-347, DOI: 10.3897/zookeys.1227.138100
EF608808121A551A880CA064B7D9A843.taxon	diagnosis	Diagnosis. Larva. The following combination of characters distinguishes P. plana from other species of Papuanatula s. str.: body dorsally without row of long, fine, simple setae along midline; body dorsally without protuberances; femur without distinct markings; abdominal segment IV with dark brown, medioposterior mark; paracercus with 9 segments.	en	Kaltenbach, Thomas, Kluge, Nikita J., Gattolliat, Jean-Luc (2025): Redescription of Papuanatula Lugo-Ortiz & McCafferty (Ephemeroptera, Baetidae), with description of a new subgenus and 20 new species. ZooKeys 1227: 159-347, DOI: 10.3897/zookeys.1227.138100
EF608808121A551A880CA064B7D9A843.taxon	description	Description. Larva (Figs 17 – 22). Body length 3.7 – 4.1 mm, cerci much longer than body length. Coloration (Fig. 17 a – c). Description see Lugo-Ortiz and McCafferty (1999: 66 – 67). Head. Antenna (Fig. 17 a). Length 1.5 × head length. As typical for the subgenus. Developing turbinate eyes in last instar male larva (Fig. 20 e) large, subquadrangular, nearly touching each other in the middle. Labrum (Fig. 18 a, b). Length 0.5 × maximum width, laterally convex. Dorsal, sub-marginal arc with ~ 14 feathered setae. Right mandible (Fig. 18 d, e). Margin between prostheca and mola with row of minute denticles. Otherwise, as typical for subgenus. Left mandible (Fig. 18 f, g). Margin between prostheca and mola with row of minute denticles. Otherwise, as typical for subgenus. Hypopharynx (Fig. 18 c). As typical for genus. Maxilla (Fig. 19 c, d). Maxillary palp slightly longer than galea-lacinia, robust; palp segment II subequal in length to segment I. Otherwise, as typical for genus. Labium (Fig. 19 a, b). As typical for the genus. Paraglossa dorsally with two spine-like setae near inner, distolateral margin. Labial palp with segment I subequal in length to segments II and III combined. Segment II with slight, broadly rounded, distomedial protuberance, dorsally with row of three spine-like setae near outer, distolateral margin. Segment III slightly pentagonal, pointed, 0.8 × length of segment II. Thorax. Sterna. With small protuberances on sides of prosternum and close to openings of mesothoracic and metathoracic sternal apodemes (as in Fig. 108 a). Terga without protuberances. Legs (Fig. 20 a – d). Ratio of leg segments: fore leg 0.9: 1.0: 0.4: 0.2, middle leg 0.9: 1.0: 0.3: 0.2 and hind leg 1.0: 1.0: 0.4: 0.2. Femur. Length ~ 3 × maximum width. Claw with one row of 5 – 8 denticles, distalmost denticle with distance to other denticles; one posterior seta. Abdomen. Terga (Figs 21 a, 22 a – d). Abdominal terga without protuberances. Posterior margin of terga: I and II smooth, without denticles; III – IX with triangular, pointed denticles, increasing in length toward VII. Surface with scattered small, sub rectangular, apically rounded scales. Tergalii (Fig. 21 e). Narrow-elongate, tracheation absent or poorly developed; margins smooth, with short, fine, simple setae. Paraproct (Fig. 21 d). Posterior margin without prolongation, smooth. Caudalii (Fig. 21 b, c). Cerci without swimming setae. Paracercus with nine segments. Pose of subimaginal gonostyli under larval cuticle (Fig. 21 f). As typical for the subgenus. Segment III conical. Subimago. Unknown. Male and female imagos. See description in Lugo-Ortiz and McCafferty (1999: 67, figs 31, 32). Imago, male (Figs 23 a, b, 24 a, b). Head and thorax yellow-brown; legs pale yellow-brown. Turbinate eyes widened apically. Fore wings marginally with double intercalary veins. Pterostigma with four mostly incomplete, oblique cross veins. Hind wings absent. Abdominal segments I – VI translucent, abdominal tergum IV posteromedially with dark brown marking similar to larvae; abdominal segments VII – X pale yellow-brown. Genitalia (Fig. 24 c). Sterno-styligeral muscle absent. Each unistyliger parallel-sided, equally wide at base and at apex. Gonostylus 1 st segment roundly-convex at apex, gradually turning to 2 nd segment. Second segment equally wide along its length. Third (terminal) segment of gonostylus nearly as wide as 2 nd segment, with length ~ 1.5 × width. Penial bridge with slightly truncated trapezoid projection between unistyligers. Egg. Unknown.	en	Kaltenbach, Thomas, Kluge, Nikita J., Gattolliat, Jean-Luc (2025): Redescription of Papuanatula Lugo-Ortiz & McCafferty (Ephemeroptera, Baetidae), with description of a new subgenus and 20 new species. ZooKeys 1227: 159-347, DOI: 10.3897/zookeys.1227.138100
EF608808121A551A880CA064B7D9A843.taxon	distribution	Distribution. New Guinea (Fig. 146).	en	Kaltenbach, Thomas, Kluge, Nikita J., Gattolliat, Jean-Luc (2025): Redescription of Papuanatula Lugo-Ortiz & McCafferty (Ephemeroptera, Baetidae), with description of a new subgenus and 20 new species. ZooKeys 1227: 159-347, DOI: 10.3897/zookeys.1227.138100
EDD2250FCAF45D02B9FDA7198F527BE3.taxon	description	Figs 114, 115, 116, 117, 118, 119, 120	en	Kaltenbach, Thomas, Kluge, Nikita J., Gattolliat, Jean-Luc (2025): Redescription of Papuanatula Lugo-Ortiz & McCafferty (Ephemeroptera, Baetidae), with description of a new subgenus and 20 new species. ZooKeys 1227: 159-347, DOI: 10.3897/zookeys.1227.138100
EDD2250FCAF45D02B9FDA7198F527BE3.taxon	etymology	Etymology. The species name pluresetae is based on the Latin words plures setae meaning “ several setae ” and refers to the three or four posterior setae on the claws.	en	Kaltenbach, Thomas, Kluge, Nikita J., Gattolliat, Jean-Luc (2025): Redescription of Papuanatula Lugo-Ortiz & McCafferty (Ephemeroptera, Baetidae), with description of a new subgenus and 20 new species. ZooKeys 1227: 159-347, DOI: 10.3897/zookeys.1227.138100
EDD2250FCAF45D02B9FDA7198F527BE3.taxon	diagnosis	Diagnosis. Larva. The following combination of characters distinguishes P. pluresetae sp. nov. from other species of Papuanatula s. str.: body dorsally with irregular row of long, fine, simple setae along midline; abdominal terga without protuberances; femur basally with short, wedge-shaped blank; otherwise, brown (after 35 years in alcohol); paracercus vestigial; claw with three or four posterior setae.	en	Kaltenbach, Thomas, Kluge, Nikita J., Gattolliat, Jean-Luc (2025): Redescription of Papuanatula Lugo-Ortiz & McCafferty (Ephemeroptera, Baetidae), with description of a new subgenus and 20 new species. ZooKeys 1227: 159-347, DOI: 10.3897/zookeys.1227.138100
EDD2250FCAF45D02B9FDA7198F527BE3.taxon	description	Description. Larva (Figs 114 – 120). Body length 4.5 – 5.8 mm, cerci much longer than body length (~ 1.3 ×). Cuticular coloration (Fig. 114 a – c; after 35 years in alcohol). Head, thorax and abdomen dorsally brown. Femur basally with short, wedge-shaped blank; otherwise, brown. Tibia pale brown; tarsus darker brown. Head, thorax, and abdomen ventrally brown. Cerci pale brown. Hypodermal coloration (Fig. 114 a, b. Abdominal terga I – IX with narrow, dark brown transverse band along posterior margins. Head. Dorsally with irregular row of long, fine, simple setae along midline (as in Fig. 26 b). Antenna (Fig. 117 e). Length ~ 1.5 × head length. Developing turbinate eyes in last instar male larva (Fig. 117 e) roundish, rather small, with large distance to each other. Labrum (Fig. 115 a, b). Length 0.5 × maximum width, laterally convex. Dorsal, sub-marginal arc with ~ 40 feathered setae. Right mandible (Fig. 115 d, e). Margin between prostheca and mola with row of minute denticles. Otherwise, as typical for subgenus. Left mandible (Fig. 115 f, g). Margin between prostheca and mola with row of minute denticles. Otherwise, as typical for subgenus. Hypopharynx (Fig. 115 c). As typical for genus. Maxilla (Fig. 116 c, d). Maxillary palp subequal in length to galea-lacinia, robust; palp segment II as long as segment I. Otherwise, as typical for genus. Labium (Fig. 116 a, b). As typical for the genus. Paraglossa dorsally with two spine-like setae near inner, distolateral margin. Labial palp with segment I 0.9 × length of segments II and III combined. Segment II with minute distomedial protuberance, dorsally with row of four spine-like setae near outer, distolateral margin. Segment III conical, pointed; 0.7 × length of segment II; inner dorsal margin with few feathered setae. Thorax. Sterna. With small protuberances on sides of prosternum and close to openings of mesothoracic and metathoracic sternal apodemes (as Fig. 108 a). Terga (Fig. 120 a) without protuberances; with irregular row of long, fine, simple setae along midline (as in Fig. 26 b). Legs (Fig. 117 a, d). Ratio of leg segments: fore leg 0.9: 1.0: 0.3: 0.2, middle leg 0.9: 1.0: 0.3: 0.2 and hind leg 1.1: 1.0: 0.4: 0.2. Femur. Length ~ 3 × maximum width. Claw with one row of seven or eight denticles and three or four posterior setae. Abdomen. Terga (Figs 119 a, 120 b – d) with irregular row of long, fine, simple setae along midline (as in Fig. 26 b). Terga without protuberances, terga I – IV with slight, paired, medioposterior elevations. Posterior margin of terga: I smooth, without denticles; II – IX with triangular, pointed denticles, partly with minute, pointed denticles in between. Surface with scattered small, triangular, pointed, striated scales. Tergalii (Fig. 118 a, b). Ovoid, tracheation poorly developed; margins smooth, with short, fine, simple setae. Paraproct (Fig. 118 c – f). Posterior margin with prolongation and row of minute denticles; on surface an area with notched scales. Caudalii (Fig. 117 f, g). Cerci apart from basal and distal part with 1 – 4 swimming setae per segment, initially increasing and then again decreasing toward distal part. Paracercus vestigial. Pose of subimaginal gonostyli under larval cuticle. As typical for the subgenus. Subimago. Unknown. Imago. Unknown. Egg. Unknown.	en	Kaltenbach, Thomas, Kluge, Nikita J., Gattolliat, Jean-Luc (2025): Redescription of Papuanatula Lugo-Ortiz & McCafferty (Ephemeroptera, Baetidae), with description of a new subgenus and 20 new species. ZooKeys 1227: 159-347, DOI: 10.3897/zookeys.1227.138100
EDD2250FCAF45D02B9FDA7198F527BE3.taxon	distribution	Distribution. New Guinea (Fig. 148).	en	Kaltenbach, Thomas, Kluge, Nikita J., Gattolliat, Jean-Luc (2025): Redescription of Papuanatula Lugo-Ortiz & McCafferty (Ephemeroptera, Baetidae), with description of a new subgenus and 20 new species. ZooKeys 1227: 159-347, DOI: 10.3897/zookeys.1227.138100
F4E3297889355B8F9D0BCA0E10E01329.taxon	description	Fig. 25 a, b	en	Kaltenbach, Thomas, Kluge, Nikita J., Gattolliat, Jean-Luc (2025): Redescription of Papuanatula Lugo-Ortiz & McCafferty (Ephemeroptera, Baetidae), with description of a new subgenus and 20 new species. ZooKeys 1227: 159-347, DOI: 10.3897/zookeys.1227.138100
F4E3297889355B8F9D0BCA0E10E01329.taxon	diagnosis	Diagnosis. Larva. The following combination of characters distinguishes P. tuber from other species of Papuanatula s. str.: body dorsally without row of long, fine, simple setae along midline; abdominal terga I – VIII (IX) with short, stout, dorsally oriented, medial protuberances; pronotum with paired, medioposterior protuberances; femur without distinct markings; paracercus vestigial; body size 2.7 – 3.4 mm.	en	Kaltenbach, Thomas, Kluge, Nikita J., Gattolliat, Jean-Luc (2025): Redescription of Papuanatula Lugo-Ortiz & McCafferty (Ephemeroptera, Baetidae), with description of a new subgenus and 20 new species. ZooKeys 1227: 159-347, DOI: 10.3897/zookeys.1227.138100
F4E3297889355B8F9D0BCA0E10E01329.taxon	description	Description. Larva (Fig. 25 a, b). See Lugo-Ortiz and McCafferty (1999: 68 – 69, figs 33 – 35). Subimago. Unknown. Imago. Unknown. Egg. Unknown.	en	Kaltenbach, Thomas, Kluge, Nikita J., Gattolliat, Jean-Luc (2025): Redescription of Papuanatula Lugo-Ortiz & McCafferty (Ephemeroptera, Baetidae), with description of a new subgenus and 20 new species. ZooKeys 1227: 159-347, DOI: 10.3897/zookeys.1227.138100
F4E3297889355B8F9D0BCA0E10E01329.taxon	distribution	Distribution. New Guinea (Fig. 146).	en	Kaltenbach, Thomas, Kluge, Nikita J., Gattolliat, Jean-Luc (2025): Redescription of Papuanatula Lugo-Ortiz & McCafferty (Ephemeroptera, Baetidae), with description of a new subgenus and 20 new species. ZooKeys 1227: 159-347, DOI: 10.3897/zookeys.1227.138100
90E494EEA5B65BCE963A9208928A618A.taxon	diagnosis	Diagnosis. Larva. The following combination of characters distinguishes P. vaisisi from other species of Papuanatula s. str.: body dorsally without row of long, fine, simple setae along midline; abdominal terga without protuberances; femur with brown, triangular mark in posterior 1 / 2; maxillary palp robust, shorter than galea-lacinia; labial palp segment III with poorly developed lateral convexity.	en	Kaltenbach, Thomas, Kluge, Nikita J., Gattolliat, Jean-Luc (2025): Redescription of Papuanatula Lugo-Ortiz & McCafferty (Ephemeroptera, Baetidae), with description of a new subgenus and 20 new species. ZooKeys 1227: 159-347, DOI: 10.3897/zookeys.1227.138100
90E494EEA5B65BCE963A9208928A618A.taxon	description	Description. Larva. See Lugo-Ortiz and McCafferty (1999: 69, figs 33 – 35) and Demoulin (1969: 229 – 230, fig. 4 a – i). Subimago. Unknown. Imago. Unknown. Egg. Unknown.	en	Kaltenbach, Thomas, Kluge, Nikita J., Gattolliat, Jean-Luc (2025): Redescription of Papuanatula Lugo-Ortiz & McCafferty (Ephemeroptera, Baetidae), with description of a new subgenus and 20 new species. ZooKeys 1227: 159-347, DOI: 10.3897/zookeys.1227.138100
90E494EEA5B65BCE963A9208928A618A.taxon	distribution	Distribution. New Britain.	en	Kaltenbach, Thomas, Kluge, Nikita J., Gattolliat, Jean-Luc (2025): Redescription of Papuanatula Lugo-Ortiz & McCafferty (Ephemeroptera, Baetidae), with description of a new subgenus and 20 new species. ZooKeys 1227: 159-347, DOI: 10.3897/zookeys.1227.138100
0C5C2BE9C0A756DF9D30E0C0C10D840B.taxon	description	Figs 121, 122, 123, 124, 125, 126	en	Kaltenbach, Thomas, Kluge, Nikita J., Gattolliat, Jean-Luc (2025): Redescription of Papuanatula Lugo-Ortiz & McCafferty (Ephemeroptera, Baetidae), with description of a new subgenus and 20 new species. ZooKeys 1227: 159-347, DOI: 10.3897/zookeys.1227.138100
0C5C2BE9C0A756DF9D30E0C0C10D840B.taxon	etymology	Etymology. The species is dedicated to Jeff Michael Webb (Rhithron Associates, USA), who was contributing to this study in an early phase.	en	Kaltenbach, Thomas, Kluge, Nikita J., Gattolliat, Jean-Luc (2025): Redescription of Papuanatula Lugo-Ortiz & McCafferty (Ephemeroptera, Baetidae), with description of a new subgenus and 20 new species. ZooKeys 1227: 159-347, DOI: 10.3897/zookeys.1227.138100
0C5C2BE9C0A756DF9D30E0C0C10D840B.taxon	diagnosis	Diagnosis. Larva. The following combination of characters distinguishes P. webbi sp. nov. from other species of Papuanatula s. str.: body dorsally without row of long, fine, simple setae along midline; pronotum with pair of small, triangular, apically rounded protuberances near medioposterior margin; fore protoptera with pair of minute protuberances at medioposterior margin; metanotum and abdominal terga I – IX posteromedially with small to medium protuberance, oriented dorsoposteriad; femur with subquadrangular blank in basal area and long, narrow blank along dorsal margin; paracercus with eight segments.	en	Kaltenbach, Thomas, Kluge, Nikita J., Gattolliat, Jean-Luc (2025): Redescription of Papuanatula Lugo-Ortiz & McCafferty (Ephemeroptera, Baetidae), with description of a new subgenus and 20 new species. ZooKeys 1227: 159-347, DOI: 10.3897/zookeys.1227.138100
0C5C2BE9C0A756DF9D30E0C0C10D840B.taxon	description	Description. Larva (Figs 121 – 126). Body length 2.4 – 3.2 mm, cerci much longer than body length (~ 1.5 ×). Cuticular coloration (Fig. 121 a – c). Head, thorax and abdomen dorsally brown. Femur grey-brown, with subquadrangular blank in basal area, long, narrow blank along dorsal margin and rounded blank in distal area; tibia and tarsus ecru, in medial area grey-brown. Head, thorax and abdomen ventrally pale brown, protuberances of thoracic sterna brown, abdominal segments II – VIII slightly darker than thorax. Cerci yellow-brown. Hypodermal coloration (Fig. 121 a, b). Abdominal terga I – IX with narrow, dark brown transverse band close to posterior margin. Head. Antenna (Fig. 121 a – c). Length 1.5 × head length. As typical for subgenus. Developing turbinate eyes in last instar male larva (Fig. 123 e) rather large, but with distance to each other. Labrum (Fig. 122 a). Relatively small, length 0.5 × maximum width, laterally slightly convex. Dorsal, sub-marginal arc with ~ 10 feathered setae. Right mandible (Fig. 122 c, d). Margin between prostheca and mola with row of minute denticles. Otherwise, as typical for subgenus. Left mandible (Fig. 122 e, f). Margin between prostheca and mola with few minute denticles. Otherwise, as typical for subgenus. Hypopharynx (Fig. 122 b). As typical for genus. Maxilla (Fig. 123 c, d). Maxillary palp subequal in length to galea-lacinia; palp segment II ~ 1.7 × length of segment I. Otherwise, as typical for genus. Labium (Fig. 123 a, b). As typical for the genus. Paraglossa dorsally with two spine-like setae near inner, distolateral margin. Labial palp with segment I ~ 1.2 × length of segments II and III combined. Segment II without distomedial protuberance, dorsally with row of four spine-like setae near outer, distolateral margin. Segment III conical, pointed, 0.7 × length of segment II. Thorax. Sterna. With small protuberances on sides of prosternum and close to openings of mesothoracic and metathoracic sternal apodemes (as in Fig. 108 a). Terga (Figs 124 f – I, 126 a). Pronotum with pair of small, triangular, apically rounded protuberances near posteromedial margin; metanotum posteromedially with small, apically rounded protuberance. Fore protoptera with pair of minute protuberances at medioposterior margin. Hind protoptera (Fig. 124 g). Absent, but sometimes with vestiges. Legs (Fig. 124 a – e). Ratio of leg segments: fore leg 0.9: 1.0: 0.3: 0.1, middle leg 0.9: 1.0: 0.3: 0.1 and hind leg 1.0: 1.0: 0.4: 0.2. Femur. Length ~ 3 × maximum width. Claw with one row of six denticles; one posterior seta. Abdomen. Terga (Figs 125 a, b, 126 b – d). Terga I – IX posteromedially with small to medium, apically rounded protuberance, oriented dorsoposteriad. Posterior margin of terga: I – III smooth, without denticles; IV – IX with short, rounded denticles, apically carrying minute, fine, acute spines. Surface with scattered small, spoon-shaped, striated scales. Tergalii (Fig. 125 f). Narrow oblong, tracheation with strongly developed trunk, other tracheation poorly developed; margins smooth, with few short, fine, simple setae. Paraproct (Fig. 125 e). Posterior margin with prolongation and row of minute denticles. Caudalii (Fig. 125 c, d). Cerci without swimming setae, in middle part one or two insertions per segment still visible, and sometimes with a short, rudimentary swimming seta. Paracercus with eight segments. Pose of subimaginal gonostyli under larval cuticle. Unknown. Subimago. Unknown. Imago. Unknown. Egg. Unknown.	en	Kaltenbach, Thomas, Kluge, Nikita J., Gattolliat, Jean-Luc (2025): Redescription of Papuanatula Lugo-Ortiz & McCafferty (Ephemeroptera, Baetidae), with description of a new subgenus and 20 new species. ZooKeys 1227: 159-347, DOI: 10.3897/zookeys.1227.138100
0C5C2BE9C0A756DF9D30E0C0C10D840B.taxon	distribution	Distribution. New Guinea (Fig. 148).	en	Kaltenbach, Thomas, Kluge, Nikita J., Gattolliat, Jean-Luc (2025): Redescription of Papuanatula Lugo-Ortiz & McCafferty (Ephemeroptera, Baetidae), with description of a new subgenus and 20 new species. ZooKeys 1227: 159-347, DOI: 10.3897/zookeys.1227.138100
94C0A0452CA0519E979CBDDEE814C6DF.taxon	description	Figs 127, 128, 129, 130, 131	en	Kaltenbach, Thomas, Kluge, Nikita J., Gattolliat, Jean-Luc (2025): Redescription of Papuanatula Lugo-Ortiz & McCafferty (Ephemeroptera, Baetidae), with description of a new subgenus and 20 new species. ZooKeys 1227: 159-347, DOI: 10.3897/zookeys.1227.138100
94C0A0452CA0519E979CBDDEE814C6DF.taxon	etymology	Etymology. The species name zebrata refers to the hypodermal coloration of larval abdomen (and probably that of winged stages) which includes contrasting, dark brown transverse bands on posterior margins of terga (Fig. 127 a, d).	en	Kaltenbach, Thomas, Kluge, Nikita J., Gattolliat, Jean-Luc (2025): Redescription of Papuanatula Lugo-Ortiz & McCafferty (Ephemeroptera, Baetidae), with description of a new subgenus and 20 new species. ZooKeys 1227: 159-347, DOI: 10.3897/zookeys.1227.138100
94C0A0452CA0519E979CBDDEE814C6DF.taxon	diagnosis	Diagnosis. Larva. The following combination of characters distinguishes P. zebrata sp. nov. from other species of Papuanatula s. str.: body without row of long, fine setae along midline; abdominal terga without protuberances; femur with brown, hypodermal streak in basal 1 / 2 and brown spot in distal area; tergalii colorless; posterior margin of abdominal terga with heterogenous, sharply pointed denticles.	en	Kaltenbach, Thomas, Kluge, Nikita J., Gattolliat, Jean-Luc (2025): Redescription of Papuanatula Lugo-Ortiz & McCafferty (Ephemeroptera, Baetidae), with description of a new subgenus and 20 new species. ZooKeys 1227: 159-347, DOI: 10.3897/zookeys.1227.138100
94C0A0452CA0519E979CBDDEE814C6DF.taxon	description	Description. Larva (Figs 127 – 130). Cuticular coloration. Head, pronotum, mesonotum and metanotum ochre with brownish areas; fore protoptera nearly uniformly ochre (Fig. 127 a). Thoracic pleura brownish, sterna mostly colorless. Cuticle of femur with brownish margins and large blank occupying most part of proximal 1 / 2 (Fig. 129 c – f). Tibia and tarsus mostly ochre (Fig. 129 c – f). Abdominal terga brownish with paler blanks; each tergum VI – IX with median blank and pair of brown sigilla inside it. Sterna mostly colorless (Fig. 127 b). Cerci uniformly pale brownish. Hypodermal coloration. Anterior side of each femur with roundish or longitudinal, dark brown macula on proximal 1 / 2; posterior side of each femur with or without brown macula on distal 1 / 2 (Fig. 129 c – f). Each abdominal tergum I – IX with contrasting, dark brown band on posterior margin; some terga also with pair of brown spots (Fig. 127 d). Tissues surrounding tracheae of tergalii either with brown pigmentation (Fig. 127 b), or without pigmentation, so that tracheae poorly visible (Fig. 127 a). Head. Antenna (Fig. 130 a). Length ~ 2 × head length. As typical for subgenus. Developing turbinate eyes in last instar male larva with facets equally developed on middle and periphery areas (as in Fig. 32 d). Labrum (Fig. 128 a) slightly widened distally; long setae on dorsal surface forming regular transverse row; each seta pointed, with moderately long processes on both sides. Right mandible (Fig. 128 e, f). Incisor with indistinct denticles near base; kinetodontium slightly separated from incisor and terminated with three denticles, with distal denticle longest. Left mandible (Fig. 128 d). Incisor and kinetodontium non-distinguishable, together with three small denticles proximad of stretched apex of incisor. Hypopharynx (Fig. 128 h) apically with pair of fields of stout, short, setae-like spines on apex. Maxilla (Fig. 128 g). Maxillary palp as long as galea-lacinia. Otherwise, as typical for genus. Labium (Fig. 128 b, c). Paraglossae widest at base and narrowing toward apex; three apical setal rows bent at apex of paraglossa. Glossa shorter than half of paraglossa, with finger-like (distal) portion as long as triangular (proximal) portion. Glossa with several long setae in distal 1 / 2 and one long seta near middle of ventral side. Labial palp without distomedian projection on segment II; segment III with median margin as long as lateral margin. Thorax. Sterna without protuberances. Terga without protuberances. Metanotum without hind protoptera or their vestiges. Legs (Fig. 129 a – f). Fore femur widened in proximal part; hind tibia shorter than others. Femur. Outer side of each femur with single regular row of long, hair-like setae bearing numerous fine, short branches on all sides (Fig. 129 c – f, as in Figs 41 g, 68 b). Tibia. Patella-tibial suture present on all legs, terminated near middle of inner margin of tibia (Fig. 129 c – f). Tibia-tarsal condylus turned to anterior side. Anterior side of each tibia with regular row of hair-like setae similar to setae on femur, but smaller (Fig. 129 c – f). Tarsus. Anterior side of each tarsus with regular row of similar, but smaller setae (Fig. 129 b). Posterior side of each tarsus with regular row of stout setae; two or three most distal of them elongated and pointed; most distal seta longer than others (but with shape and thickness similar to previous one) (Fig. 129 a). Claw with row of 4 – 6 denticles and one somewhat larger denticle distad of them; long, arched, posterior seta (Fig. 129 b). Abdomen. Terga (Figs 127 b, 130 b, c) without long setae on midline. Abdominal terga without dorsal unpaired or paired protuberances, only with slightly expressed, unpaired, median elevations. Abdominal terga I – III without denticles or with few denticles on posterior margins; posterior margins of abdominal terga IV – IX with conical, sharply pointed denticles irregularly alternated with smaller pointed denticles. Posterior margin of tergum X mostly smooth, with few pointed denticles on sides. Tergalii (Fig. 127 a, b) of abdominal segment I absent; tergalii II – VII subequal, oval. Each tergalius with costal and anal ribs narrow, smooth, present on proximal 1 / 2 of tergalius only (as in Fig. 66 i). Paraproct without posterior prolongation; margins membranous, smooth, lacking denticles. Caudalii (Fig. 127 a, b) without swimming setae or their vestiges. Paracercus short, consisting of ~ 8 – 10 segments. Pose of subimaginal gonostyli under larval cuticle (Fig. 131 b). In mature larva ready to molt to subimago, subimaginal gonostyli packed under larval cuticle in “ Labiobaetis - type ” pose: 2 nd segments directed medially and bent proximally; 3 rd segment directed medially (as continuation of 2 nd segment) and narrowed apically, being deformed corresponding to space between subimaginal styliger and larval cuticle. Subimago. Cuticular coloration. Pronotum and mesonotum on Fig. 131 a. Texture. On all legs of both sexes, each tarsomere covered mostly with blunt microlepides, with pointed microlepides near apex (as in Fig. 70 i). Imago. Unknown. Judging from larval hypodermal coloration and identical hypodermal coloration of male and female larvae ready to molt to subimago, imagines of both sexes have following features: Femur of each leg pair with two dark brown maculae, one on proximal 1 / 2 and another on distal 1 / 2. Each abdominal tergum I – IX with contrasting, dark brown band on posterior margin; some terga also with pair of brown spots (as in Fig. 127 d). Judging from fully developed facetted surface of turbinate eyes in larva, male imaginal turbinate eyes are widened distally and have wide facetted surfaces. Egg. Unknown.	en	Kaltenbach, Thomas, Kluge, Nikita J., Gattolliat, Jean-Luc (2025): Redescription of Papuanatula Lugo-Ortiz & McCafferty (Ephemeroptera, Baetidae), with description of a new subgenus and 20 new species. ZooKeys 1227: 159-347, DOI: 10.3897/zookeys.1227.138100
94C0A0452CA0519E979CBDDEE814C6DF.taxon	distribution	Distribution. New Guinea (Fig. 148).	en	Kaltenbach, Thomas, Kluge, Nikita J., Gattolliat, Jean-Luc (2025): Redescription of Papuanatula Lugo-Ortiz & McCafferty (Ephemeroptera, Baetidae), with description of a new subgenus and 20 new species. ZooKeys 1227: 159-347, DOI: 10.3897/zookeys.1227.138100
67323A685CF85B53A343ED508B4AE6DF.taxon	type_taxon	Type species. Papuanatula tuber Lugo-Ortiz & McCafferty, 1999, by original designation (Lugo-Ortiz and McCafferty 1999)	en	Kaltenbach, Thomas, Kluge, Nikita J., Gattolliat, Jean-Luc (2025): Redescription of Papuanatula Lugo-Ortiz & McCafferty (Ephemeroptera, Baetidae), with description of a new subgenus and 20 new species. ZooKeys 1227: 159-347, DOI: 10.3897/zookeys.1227.138100
67323A685CF85B53A343ED508B4AE6DF.taxon	diagnosis	Diagnosis. Larval characters. The following combination of characters distinguishes Papuanatula from all other genera of Baetidae: antennal scapes basally narrow, distally broad; labrum much wider than long; labium with glossae much shorter than paraglossae; maxillary palp with two segments; both mandibles outside laterally with some long, robust, simple setae; outer side of each femur usually with single regular row of long, hair-like setae; femoral patch absent on all legs; patella-tibial suture usually present on all legs; tibia-tarsal condylus (originally located on outer side) turned to anterior side; anterior side of each tibia usually with regular row of setae similar to that on femur; anterior side of each tarsus usually with regular row of setae similar to femur and tibia; tarsus with conspicuous, long seta subdistally on inner margin; claw with single row of denticles and one or several posterior setae; hind protoptera absent or vestigial; tergalii present on abdominal segments II – VII; paraproct usually with prolongation on proximal margin; cerci usually longer than body length; paracercus strongly reduced or vestigial.	en	Kaltenbach, Thomas, Kluge, Nikita J., Gattolliat, Jean-Luc (2025): Redescription of Papuanatula Lugo-Ortiz & McCafferty (Ephemeroptera, Baetidae), with description of a new subgenus and 20 new species. ZooKeys 1227: 159-347, DOI: 10.3897/zookeys.1227.138100
67323A685CF85B53A343ED508B4AE6DF.taxon	description	Description. Male imago. Fore wings. With marginal double intercalary veins. Hind wings absent. Genitalia. Sterno-styliger muscle completely absent. Gonostyli segment II without significant widenings. Segment III (terminal) of gonostyli nearly as wide as segment II, length varying from slightly exceeding width to twice width.	en	Kaltenbach, Thomas, Kluge, Nikita J., Gattolliat, Jean-Luc (2025): Redescription of Papuanatula Lugo-Ortiz & McCafferty (Ephemeroptera, Baetidae), with description of a new subgenus and 20 new species. ZooKeys 1227: 159-347, DOI: 10.3897/zookeys.1227.138100
67323A685CF85B53A343ED508B4AE6DF.taxon	distribution	Distribution. Papua New Guinea, Indonesia: New Guinea, Sulawesi.	en	Kaltenbach, Thomas, Kluge, Nikita J., Gattolliat, Jean-Luc (2025): Redescription of Papuanatula Lugo-Ortiz & McCafferty (Ephemeroptera, Baetidae), with description of a new subgenus and 20 new species. ZooKeys 1227: 159-347, DOI: 10.3897/zookeys.1227.138100
