Papuanatula (Papuanatula) bessa Lugo-Ortiz & McCafferty, 1999

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, pp. 159-347 : 159-347

publication ID	https://doi.org/10.3897/zookeys.1227.138100
publication LSID	lsid:zoobank.org:pub:F259B26F-4DA1-452E-ABEE-7D0957CFE261
DOI	https://doi.org/10.5281/zenodo.14852556
persistent identifier	https://treatment.plazi.org/id/3EF77030-6D6A-5AF7-B1A5-9EAC62EA7461
treatment provided by	ZooKeys by Pensoft
scientific name	Papuanatula (Papuanatula) bessa Lugo-Ortiz & McCafferty, 1999
status

Treatment

Papuanatula (Papuanatula) bessa Lugo-Ortiz & McCafferty, 1999 View in CoL

Figs 1 View Figure 1 , 2 View Figure 2 , 3 View Figure 3 , 4 View Figure 4 , 5 View Figure 5 , 6 View Figure 6

Papuanatula bessa Lugo-Ortis & McCafferty, 1999: 63 View in CoL (larva, ♂ and ♀ subimago).

Material examined.

Type locality (‘ additional material’ of original description). Papua New Guinea • 15 larvae; Morobe Prov., E of Wau, Bulolo Riv. ; 900 m; 18. x. 1964; leg. WL and WG Peters; 3 on slides; GBIFCH 00592634 , GBIFCH 00976069 , GBIFCH 01221771 ; MZL; 12 in alcohol; GBIFCH 00976073 , GBIFCH 00975821 ; MZL .

Other material.

Papua New Guinea • 6 larvae; Gulf Prov., Marawaka, Mala ; 07°05'40"S, 145°44'28"E; 1400 m; 11. xi. 2006; leg. M. Balke & Kinibel; (PNG 90); 1 on slide; GBIFCH 00976130 ; MZL; 5 in alcohol; GBIFCH 00975772 , GBIFCH 00976022 , GBIFCH 00976129 , GBIFCH 00976131 ; MZL GoogleMaps • 8 larvae; Morobe Prov., Menyamya, Mt Inji ; near 07°14'49"S, 146°01'20"E; 1700 m; 14. xi. 2006; leg. M. Balke & Kinibel; (PNG 96); in alcohol; GBIFCH 00975779 , GBIFCH 00976085 ; MZL GoogleMaps • 2 larvae; Central Prov., Tapini ; 08°20'31"S, 146°59'49"E; 870 m; 29. x. 2007; leg. Kinibel; (PNG 161); in alcohol; GBIFCH 00975785 , GBIFCH 00976094 ; MZL GoogleMaps • 9 larvae; Morobe Prov., Wagau, Herzog Mts ; 06°51'04"S, 146°48'04"E; 1150 m; 19. xi. 2006; leg. M. Balke & Kinibel; (PNG 102); 1 on slide; GBIFCH 00592633 ; MZL; 8 in alcohol; GBIFCH 00975777 , GBIFCH 00976081 , GBIFCH 00976082 , GBIFCH 00976083 ; MZL GoogleMaps . INDONESIA • Papua, Waena ; 8–13. viii. 2012; coll. N. Kluge & L. Sheyko: 3 L-S ♂, 13 larvae; SPbU .

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 View Figure 1 ; Lugo-Ortiz and McCafferty 1999: fig. 5); femur brownish, with wedge-shaped blank in proximal 1 / 2; paracercus with eight segments.

Description.

Larva (Figs 1 a View Figure 1 – 5 h View Figure 5 ). 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 View Figure 2 ). 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 View Figure 2 , 4 a View Figure 4 ). Tibia and tarsus brownish (Fig. 2 d – f View Figure 2 ). Abdominal terga mostly brownish, with lateral areas paler; terga V – VI more or less paler than others; sterna mostly colorless (Figs 1 a View Figure 1 , 2 g View Figure 2 ). Cerci uniformly pale brownish.

Hypodermal coloration. Legs without hypodermal markings. Abdominal terga with extensive, contrasting, dark brown markings of characteristic shape (Figs 1 a View Figure 1 , 2 a View Figure 2 ). 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 View Figure 26 , 65 b View Figure 65 ). Antenna (Fig. 2 a, b View Figure 2 ). 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 View Figure 66 ). Labrum (Fig. 3 a, b View Figure 3 ) 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 View Figure 3 ). 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 View Figure 3 ). 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 View Figure 3 ). As typical for genus. Maxilla (Fig. 3 g View Figure 3 ). Maxillary palp as long as galea-lacinia. Otherwise, as typical for genus. Labium (Fig. 3 c View Figure 3 ). 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 View Figure 108 ). Terga with irregular row of long, fine, soft, colorless, simple setae along midline (as in Figs 26 b View Figure 26 , 65 b View Figure 65 ). Metanotum with medioposterior broad, paired elevations. Metanotum without hind protoptera or their vestiges. Legs (Figs 2 d – f View Figure 2 , 4 a – e View Figure 4 ). 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 View Figure 41 , 68 b View Figure 68 ). 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 View Figure 5 ). Long, fine, soft, colorless setae irregularly situated along midline of all abdominal terga (as in Figs 26 b View Figure 26 , 65 b View Figure 65 ). 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 View Figure 2 , 4 f, g View Figure 4 ) 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 View Figure 5 ) with posterior prolongation bent toward bases of caudalii, with many small, equal denticles on median-posterior margin. Caudalii (Fig. 4 h, i View Figure 4 ) 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 View Figure 6 ).

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 View Figure 70 ).

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 View Figure 6 ).

Imago, female. Unknown. Judging by description of subimago ( Lugo-Ortiz and McCafferty 1999: 64), hypodermal coloration of abdominal terga as in male.

Egg. Unknown.

Dimension.

Fore wing length (and approximate body length): 4 mm.

Distribution.

New Guinea (Fig. 146 View Figure 146 ).