Research Article |
Thomas Kaltenbach‡§, Jean-Luc Gattolliat‡§
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Corresponding author: Thomas Kaltenbach ( thomas.kaltenbach@bluewin.ch ) Academic editor: Eduardo Dominguez
© 2021 Thomas Kaltenbach, Jean-Luc Gattolliat.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Kaltenbach T, Gattolliat J-L (2021) New species of Labiobaetis Novikova & Kluge from Southeast Asia and New Guinea (Ephemeroptera, Baetidae). ZooKeys 1067: 159-208. https://doi.org/10.3897/zookeys.1067.72251
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Abstract
Material collected between 2006 and 2016 in Borneo, Sulawesi, and New Guinea further increased our knowledge of Labiobaetis Novikova & Kluge in these regions. Five species were previously reported from Borneo, two from Sulawesi, and 33 from New Guinea. Six new species have been identified using a combination of morphology and genetic distance (COI, Kimura 2-parameter), one species from Borneo (Brunei), one from Sulawesi, and four from New Guinea. They are described and illustrated based on their larvae and keys to the species of the relevant groups are provided. Additionally, new reports, a complementary description, and the COI sequence for L. dendrisetis Kaltenbach & Gattolliat are presented. The distribution of Labiobaetis in the Wallacea region is discussed based on the new findings. The total number of Labiobaetis species worldwide is augmented to 153.
Keywords
Biogeography, Borneo, COI, integrated taxonomy, morphology, Sulawesi
Introduction
The family Baetidae has the highest species diversity among mayflies, comprising ca. 1,100 species in 114 genera (updated from
The genus Labiobaetis Novikova & Kluge, 1987 (
This contribution will focus on further new species of Labiobaetis from Borneo, Sulawesi and New Guinea with integrative taxonomy. In the past, five species were reported from Indonesia (L. fulmeki (Ulmer), L. obscurum (Ulmer), L. necopinatum (Müller-Liebenau), L. ulmeri (Müller-Liebenau) and L. boettgeri (Ulmer)). All were described from adults only and no species were previously known at the larval stage (
Indonesia is an immense archipelago of more than 18.000 islands extending over a huge area from 95°E to 141°E and from 6°N to 11°S. It is one of the most biologically rich countries in the world. The high levels of species richness and endemism are mainly attributable to a complex geological history, that brought together two different biological realms (Oriental realm and Australasian realm), separated by a transitional region (Wallacea) (
Materials and methods
Part of the material was collected during a series of university training practicals (see also
All specimens were preserved in 70%–96% ethanol. The dissection of larvae was done in Cellosolve (2-Ethoxyethanol) with subsequent mounting on slides with Euparal liquid, using an Olympus SZX7 stereomicroscope.
The DNA of part of the specimens was extracted using non-destructive methods allowing subsequent morphological analysis (see
The GenBank accession numbers are given in Table 1; the nomenclature of gene sequences follows
Sequenced specimens: treated species and known species of group claudiae.
| Species | Species group | Locality | Specimens catalog # | GenBank # (COI) | GenSeq Nomenclature |
|---|---|---|---|---|---|
| L. catadupa sp. nov. | catadupa | Brunei | GBIFCH00592439 | MW868314 | genseq-2 COI |
| L. toraja sp. nov. | catadupa | Sulawesi | GBIFCH00674627 | MW868315 | genseq-2 COI |
| GBIFCH00674628 | MW868316 | genseq-2 COI | |||
| L. academicus | claudiae | Papua Province | GBIFCH00673069 | MW041241 | genseq-2 COI |
| GBIFCH00673081 | MW041242 | genseq-2 COI | |||
| L. centralensis | claudiae | Papua New Guinea: Central Prov. | GBIFCH00465215 | MH619495 | genseq-1-COI |
| GBIFCH00465216 | MH619494 | genseq-2 COI | |||
| L. claudiae | claudiae | Papua New Guinea: Madang Prov. | GBIFCH00508144 | MH619479 | genseq-1-COI |
| L. hattam sp. nov. | claudiae | Papua Barat | GBIFCH00763707 | MW868311 | genseq-2 COI |
| L. stagnum | claudiae | Papua Province | GBIFCH00465168 | MH619491 | genseq-2 COI |
| L. werneri sp. nov. | claudiae | Papua New Guinea: Gulf Prov. | GBIFCH00763699 | MW868307 | genseq-1-COI |
| Papua New Guinea: Eastern Highlands | GBIFCH00763603 | MW868309 | genseq-2 COI | ||
| Papua New Guinea: Morobe Prov. | GBIFCH00763700 | MW868308 | genseq-2 COI | ||
| L. dendrisetis | dendrisetis | Papua New Guinea: Central Prov. | GBIFCH00763706 | MW868310 | genseq-4 COI |
| L. arfak sp. nov. | seramensis | Papua Barat | GBIFCH00763714 | MW868312 | genseq-2 COI |
| GBIFCH00763715 | MW868313 | genseq-2 COI |
Drawings were made using an Olympus BX43 microscope. To facilitate the determination of species and the comparison of important structures, we partly used a combination of dorsal and ventral aspects in one drawing. Explanations are given in
Photographs of larvae were taken using a Canon EOS 6D camera and processed with the programs Adobe Photoshop Lightroom (http://www.adobe.com) and Helicon Focus version 5.3 (http://www.heliconsoft.com). Photographs were subsequently enhanced with Adobe Photoshop Elements 13.
The distribution maps were generated with the program SimpleMappr (https://simplemappr.net,
The dichotomous keys were elaborated with the support of the program DKey version 1.3.0 (http://drawwing.org/dkey,
The terminology follows
Abbreviations
Results
List of Labiobaetis species treated in this paper
catadupa group (new group)
1. L. catadupa sp. nov.
2. L. toraja sp. nov.
claudiae group
3. L. hattam sp. nov.
4. L. werneri sp. nov.
dendrisetis group
5. L. dendrisetis Kaltenbach & Gattolliat, 2018
seramensis group
6. L. arfak sp. nov.
7. L. onim sp. nov.
Labiobaetis catadupa group of species (new group of species)
The catadupa group can be recognised by the following combination of characters: A) dorsal surface of labrum with submarginal arc of feathered setae (Figs 1b, 6a, 3); B) labial palp segment II extended thumb-like, glossae much shorter than paraglossae (Figs 1i, 6h); C) claws with long subapical seta on posterior side and reduced subapical seta on anterior side (Figs 2b, 4a, b, 7d); D) hind protoptera absent; E) six pairs of gills (gill I absent).
The L. catadupa group is so far known from Borneo and Sulawesi, it includes the following species:
Labiobaetis catadupa sp. nov.
Labiobaetis toraja sp. nov.
Labiobaetis catadupa sp. nov.
Type material
Holotype. Brunei • larva; Temburong District, Ulu Temburong National Park, Belalong River (near field station); 04°32'49"N, 115°09'30"E; 100 m; v. 2014; leg. K. Baker; on slide; GBIFCH00592448;
Diagnosis
Larva. Following combination of characters: A) dorsal surface of labrum with submarginal arc of 17–19 long, feathered setae with broad middle part (Figs 1a, b, 3); B) labial palp segment II with extended thumb-like distomedial protuberance, segment III slightly pentagonal; glossae much shorter than paraglossae (Fig. 1i); C) left mandible without setae at apex of mola; D) fore femur length ca. 3× maximum width, dorsal margin with 17–19 curved, spine-like setae (Fig. 2a); E) claw with long subapical seta on posterior side and reduced subapical seta on anterior side (Figs 2b, 4a); F) hind protoptera absent; G) six pairs of gills (gill I absent); H) paraproct distally expanded, with ca. 40 stout, marginal spines.
Description
Larva (Figs 1–3, 4a, 17a). Body length 2.6–6.0 mm. Cerci: ca. 2/3 of body length. Paracercus: ca. 1/5 of cerci length. Antenna: approx. twice as long as head length.
Colouration (Fig. 17a). Head, thorax and abdomen dorsally and ventrally brown. Legs light brown, caudalii light brown.
Antenna (Fig. 2f) with scape and pedicel sub cylindrical, without distolateral process at scape.
Labrum (Fig. 1a, b). Rectangular, length 0.6× maximum width. Distal margin with medial emargination and a small process. Dorsally with medium, fine, simple setae scattered over surface; submarginal arc of setae composed of 17–19 long, feathered setae with broad middle part. Ventrally with marginal row of setae composed of anterolateral long, feathered setae and medial long, bifid setae; ventral surface with ca. two short, spine-like setae near lateral and anterolateral margin.
Right mandible (Fig. 1c, d). Incisor and kinetodontium fused. Incisor with five denticles; kinetodontium with three denticles, inner margin of innermost denticle with a row of thin setae. Prostheca robust, apically denticulate. Margin between prostheca and mola slightly convex, with few minute denticles. Tuft of setae at apex of mola present.
Left mandible (Fig. 1e, f). Incisor and kinetodontium fused. Incisor with five denticles; kinetodontium with three denticles. Prostheca robust, apically with small denticles and comb-shaped structure. Margin between prostheca and mola straight, with minute denticles. Subtriangular process long and slender, above level of area between prostheca and mola. Denticles of mola apically constricted. Tuft of setae at apex of mola absent.
Both mandibles with lateral margins slightly convex. Basal half with fine, simple setae scattered over dorsal surface.
Hypopharynx and superlinguae (Fig. 1g). Lingua shorter than superlingua. Lingua approx. as long as broad; distal half laterally not expanded; medial tuft of stout setae well developed. Superlinguae distally rounded; lateral margins rounded; fine, long, simple setae along distal margin.
Maxilla (Fig. 1h). Galea-lacinia ventrally with two simple, apical setae under canines. Inner dorsal row of setae with three denti-setae, distal denti-seta tooth-like, middle and proximal denti-setae slender, bifid and pectinate. Medially with one pectinate, spine-like seta and two or three medium, simple setae. Maxillary palp ca. 1.2× length of galea-lacinia; 2-segmented; palp segment II approx. as long as segment I; setae on maxillary palp fine, simple, scattered over surface of segments I and II; apex of last segment with slight excavation at inner distolateral margin.
Labium (Fig. 1i). Glossa basally broad, narrowing toward apex; much shorter than paraglossa; inner margin with one spine-like seta; apex with two long and one medium, robust setae; outer margin with three spine-like setae; ventral surface with fine, simple, scattered setae. Paraglossa sub-rectangular, slightly curved inward; apex slightly concave; with three rows of long, robust, distally pectinate setae in apical area and two or three short, simple setae in anteromedial area; dorsally with two long, spine-like setae near inner margin. Labial palp with segment I 0.7× length of segments II and III combined. Segment I ventrally with short, fine, simple setae. Segment II with extended thumb-like, distomedial protuberance, bent upwards; distomedial protuberance 0.8× width of base of segment III; ventral surface with short, fine, simple setae; dorsally without spine-like setae near outer margin. Segment III slightly pentagonal; length 1.3× width; ventrally covered with short, spine-like, simple setae and short, fine, simple setae.
Hind protoptera (Fig. 2g) absent.
Foreleg (Fig. 2a, b). Ratio of foreleg segments 1.2:1.0:0.5:0.2. Femur. Length ca. 3× maximum width. Dorsal margin with a row of 17–19 curved, spine-like setae and a row of long, fine, simple setae; length of setae 0.24× maximum width of femur. Apex rounded, with a pair of spine-like setae and some short, stout setae. Stout, apically rounded setae scattered along ventral margin; femoral patch absent. Tibia. Dorsal margin with a row of short, spine-like setae and long, fine, simple setae. Ventral margin with a row of short, curved, spine-like setae, on apex a tuft of fine, simple setae. Patellotibial suture present on basal 1/2. Tarsus. Dorsal margin with a row of fine, simple setae. Ventral margin with a row of curved, spine-like setae. Claw with one row of eight or nine denticles; distally pointed; with ca. four stripes; with long, subapical seta on posterior side and reduced, subapical seta on anterior side.
Terga (Fig. 2c). Surface with irregular rows of U-shaped scale bases and scattered fine, simple setae. Posterior margin of tergum IV with rounded spines, wider than long.
Gills (Fig. 2d). Present on segments II–VII. Margin with small denticles intercalating fine simple setae. Tracheae extending from main trunk to inner and outer margins. Gill IV as long as length of segments V and 1/3 VI combined. Gill VII as long as length of segment VIII.
Paraproct (Fig. 2e). Distally expanded, with ca. 40 stout, marginal spines. Surface scattered with fine, simple setae. Cercotractor with numerous small, marginal spines.
Etymology
Based on the Latin word catadupa, meaning waterfall, with reference to the habitat of the species.
Distribution
Brunei (Fig. 21b).
Biological aspects
The specimens were collected at an altitude of 150 m, mostly from waterfalls with slope angles of 16° to 50° and lengths between 5 m and 20 m (Fig. 5;
Labiobaetis toraja sp. nov.
Type material
Holotype. Indonesia • larva; Sulawesi; Tengah, Lake Lore; 01°19'35"S, 120°18'40"E; 1600 m; 01.ix.2011; leg. Sumoked (SUL013); on slide; GBIFCH00592443;
Diagnosis
Larva. Following combination of characters: A) dorsal surface of labrum with submarginal arc of one plus 18–21 long, feathered setae (Figs 6a, 4b); B) labial palp segment II with hook-like distomedial protuberance, segment III oblong; glossae much shorter than paraglossae (Fig. 6h); C) left mandible with setae at apex of mola; D) fore femur length ca. 3× maximum width, dorsal margin with 18–25 curved, spine-like setae (Fig. 7a); E) claw with long subapical seta on posterior side and reduced subapical seta on anterior side (Figs 4b, 7d); F) hind protoptera absent; G) six pairs of gills (gill I absent); H) paraproct distally slightly expanded, with more than 40 stout, marginal spines.
Description
Larva (Figs 4b, 6, 7, 17b, c). Body length 5.5–6.5 mm. Cerci broken. Paracercus: ca. 0.4× body length. Antenna: approx. twice as long as head length.
Colouration (Fig. 17b, c). Head, thorax and abdomen dorsally brown, with pattern as in Fig. 17b. Head, thorax and abdomen ventrally light brown, abdominal segments VII–IX laterally darker (Fig. 17c). Legs light brown; femur with dorsomedial brown streak and brown sections apically and distoventrally; tibia basally and tarsus distally darker (Fig. 17c). Caudalii ecru.
Antenna (Fig. 7h) with scape and pedicel sub cylindrical, without distolateral process at scape. Scape and pedicel with few stout setae.
Labrum (Fig. 6a). Sub-rectangular, length 0.7× maximum width. Distal margin with medial emargination and a small process. Dorsally with medium, fine, simple setae scattered over surface; submarginal arc of setae composed of one plus 18–21 long, feathered setae. Ventrally with marginal row of setae composed of lateral and anterolateral long, feathered setae and medial long, bifid setae; ventral surface with ca. seven short, spine-like setae near lateral and anterolateral margin.
Right mandible (Fig. 6b, c). Incisor and kinetodontium fused. Incisor with five denticles; kinetodontium with three denticles, inner margin of innermost denticle with a row of thin setae. Prostheca robust, apically denticulate. Margin between prostheca and mola straight. Tuft of setae at apex of mola present.
Left mandible (Fig. 6d, e). Incisor and kinetodontium fused. Incisor with five denticles; kinetodontium with three denticles. Prostheca robust, apically with small denticles and comb-shaped structure. Margin between prostheca and mola straight, with few minute denticles. Subtriangular process long and slender, above level of area between prostheca and mola. Denticles of mola apically constricted. Tuft of setae at apex of mola present.
Both mandibles with lateral margins slightly convex. Basal half with fine, simple setae scattered over dorsal surface.
Hypopharynx and superlinguae (Fig. 6f). Lingua shorter than superlingua. Lingua approx. as long as broad; distal half laterally not expanded; medial tuft of stout setae well developed and long. Superlinguae distally rounded; lateral margins rounded; fine, long, simple setae along distal margin.
Maxilla (Fig. 6g). Galea-lacinia ventrally with five simple, apical setae under canines. Inner dorsal row of setae with three denti-setae, distal denti-seta tooth-like, middle and proximal denti-setae slender, bifid and pectinate. Medially with one pectinate, spine-like seta and 6–8 long, simple setae. Maxillary palp ca. 1.3× length of galea-lacinia; 2-segmented; palp segment II 1.2× length of segment I; setae on maxillary palp fine, simple, scattered over surface of segments I and II; apex of last segment without excavation at inner distolateral margin.
Labium (Fig. 6h, i). Glossa basally broad, narrowing toward apex; much shorter than paraglossa; inner margin with two spine-like setae; apex with two long and one medium, robust setae; outer margin with three spine-like setae; ventral surface with fine, simple, scattered setae. Paraglossa broad, slightly curved inward; outer margin convex; apex rounded; with three long rows of long, robust, distally pectinate setae in apical area, five or six short, simple setae in anteromedial area and one short, simple seta in posteromedial area; dorsally with a row of four long, spine-like setae near inner margin. Labial palp with segment I 0.8× length of segments II and III combined. Segment I ventrally with short, fine, simple setae. Segment II with hook-like, distomedial protuberance; distomedial protuberance 0.9× width of base of segment III; ventral surface with short, simple setae; dorsally with one or two spine-like setae near outer margin. Segment III oblong; length 1.7× width; ventrally covered with short, spine-like, simple setae and short, fine, simple setae.
Hind protoptera (Fig. 7i) absent.
Foreleg (Fig. 7a–d). Ratio of foreleg segments 1.2:1.0:0.6:0.2. Femur. Length ca. 3× maximum width. Dorsal margin with a row of 18–25 curved, spine-like setae and a row of long, fine, simple setae; length of setae 0.21× maximum width of femur. Apex rounded, with a pair of spine-like setae and some short, stout setae. Stout, apically rounded setae scattered along ventral margin; femoral patch absent. Tibia. Dorsal margin with a row of short, spine-like setae. Ventral margin with a row of short, curved, spine-like setae, on apex some longer setae and a tuft of fine, simple setae. Anterior surface scattered with stout, lanceolate setae. Patellotibial suture present on basal 2/3 area. Tarsus. Dorsal margin with a row of short, spine-like setae. Ventral margin with a row of curved, spine-like setae and a row of short, stout setae near margin. Claw with one row of nine or ten denticles; distally pointed; with ca. four stripes; with long, subapical seta on posterior side and reduced, subapical seta on anterior side.
Middle and hind legs. As foreleg, but middle leg with a reduced femoral patch and hind leg with a rather well developed femoral patch.
Terga (Fig. 7e). Surface with irregular rows of U-shaped scale bases. Posterior margin of tergum IV with triangular, apically rounded spines, slightly longer than wide, and fine simple setae.
Gills (Fig. 7f). Present on segments II–VII. Margin with small denticles intercalating fine simple setae. Tracheae extending from main trunk to inner and outer margins. Gill IV as long as length of segments V and 1/2 VI combined. Gill VII as long as length of segment VIII.
Paraproct (Fig. 7g). Distally slightly expanded, with more than 40 stout, marginal spines. Surface scattered with U-shaped scale bases, fine, simple setae and micropores. Cercotractor with numerous small, marginal spines.
Etymology
Dedicated to the indigenous Toraja people of Sulawesi, where the type locality is located.
Distribution
Indonesia: Sulawesi (Fig. 21b).
Biological aspects
The specimens were collected at an altitude of 1600 m in a tributary to Lake Lore.
Labiobaetis claudiae group of species (Kaltenbach and Gattolliat 2018 )
The claudiae group is recognised by the following combination of characters: A) dorsal surface of labrum with submarginal arc of simple setae; B) labial palp segment II with rather narrow thumb-like distomedial protuberance; C) maxillary palp segment II without distolateral excavation, apex usually constricted; D) six pairs of gills (gill I absent); E) gills margin usually with both shorter and longer setae; F) hind protoptera absent; G) distolateral process at scape absent; H) femur dorsally with relatively short setae (length below 0.20× maximum width of femur); I) femur apically with stout setae on posterior side of foreleg and middle leg; J) femoral patch present on all legs.
The L. claudiae group is known from New Guinea only, it includes the following species:
Labiobaetis academicus Kaltenbach, Surbakti & Kluge, 2021
Labiobaetis centralensis Kaltenbach & Gattolliat, 2018 (new assignment, see discussion)
Labiobaetis claudiae Kaltenbach & Gattolliat, 2018
Labiobaetis hattam sp. nov.
Labiobaetis stagnum Kaltenbach & Gattolliat, 2018
Labiobaetis werneri sp. nov.
Key to the species of the Labiobaetis claudiae group (larvae)
| 1 | Paraproct distally expanded (Fig. 9g) | 2 |
| – | Paraproct distally not expanded (Fig. 11f) | 4 |
| 2 | Anal margins of gills with both longer and shorter setae ( |
L. stagnum |
| – | Anal margin of gills with short setae only (Fig. 9f). | 3 |
| 3 | Posterior margins of tergites with triangular spines, longer than wide (Fig. 9e); scape with stout setae (Fig. 9h). | L. hattam sp. nov. |
| – | Posterior margins of tergites with triangular spines, wider than long ( |
L. centralensis |
| 4 | Posterior margins of tergites with triangular spines, wider than long (Fig. 11d) | L. werneri sp. nov. |
| – | Posterior margins of tergites with triangular spines, longer than wide ( |
5 |
| 5 | Labial palp segment II with rather broad thumb-like distomedial protuberance ( |
L. claudiae |
| – | Labial palp segment II with narrow thumb-like distomedial protuberance ( |
L. academicus |
Labiobaetis hattam sp. nov.
Type material
Holotype. Indonesia • larva; Papua Barat, Fumato to Kebar, forest stream; 00°52'29"S, 132°46'06"E; 492 m; 06.xi.2013; leg. UNIPA team; BH030; on slide; GBIFCH00592775;
Diagnosis
Larva. Following combination of characters: A) dorsal surface of labrum with submarginal arc of one plus six long, simple setae (Fig. 8a); B) labial palp segment II with narrow, extended, distomedial protuberance, segment III slightly pentagonal (Fig. 8h); C) fore femur rather slender, length ca. 4× maximum width, dorsal margin with 19–23 spine-like setae (Fig. 9a); D) hind protoptera absent; E) six pairs of gills (gill I absent), margin with short setae only; F) paraproct distally slightly expanded, with 38–48 stout, marginal spines (Fig. 9g); G) Scape apically with stout setae (Fig. 9h).
Description
Larva (Figs 8, 9, 18a, b). Body length 7.0–7.6 mm. Cerci and paracercus broken. Antenna: approx. 3× as long as head length.
Colouration (Fig. 18a, b). Head, thorax and abdomen dorsally dark brown, with pattern as in Fig. 18a; fore protoptera brown with bright stripes. Thorax ventrally ecru, abdomen ventrally brown, with pattern as in Fig. 18b. Legs light brown; femur with dorsomedial and apical brown spots; tibia medially and tarsus proximally brown. Caudalii brown.
Antenna (Fig. 9h) with scape and pedicel sub cylindrical, without distolateral process at scape. Scape apically with few stout setae.
Labrum (Fig. 8a). Sub-rectangular, length 0.8× maximum width. Distal margin with medial emargination and a small process. Dorsally with medium, fine, simple setae scattered over surface; submarginal arc of setae composed of one plus six long, simple setae. Ventrally with marginal row of setae composed of anterolateral long, feathered setae and medial long, bifid, pectinate setae; ventral surface with ca. seven short, spine-like setae near lateral and anterolateral margin.
Right mandible (Fig. 8b, c). Incisor and kinetodontium fused. Incisor with five denticles; kinetodontium with three denticles, inner margin of innermost denticle with a row of thin setae. Prostheca robust, apically denticulate. Margin between prostheca and mola almost straight. Tuft of setae at apex of mola present.
Left mandible (Fig. 8d, e). Incisor and kinetodontium fused. Incisor with four denticles; kinetodontium with three denticles. Prostheca robust, apically with small denticles and comb-shaped structure. Margin between prostheca and mola almost straight. Subtriangular process long and slender, above level of area between prostheca and mola. Denticles of mola apically constricted. Tuft of setae at apex of mola present.
Both mandibles with lateral margins almost straight. Basal half with fine, simple setae scattered over dorsal surface.
Hypopharynx and superlinguae (Fig. 8f). Lingua approx. as long as superlingua. Lingua longer than broad; distal half laterally slightly expanded; medial tuft of stout setae well developed and short. Superlinguae distally rounded; lateral margins rounded; fine, long, simple setae along distal margin.
Maxilla (Fig. 8g). Galea-lacinia ventrally with two simple, apical setae under canines. Inner dorsal row of setae with three denti-setae, distal denti-seta tooth-like, middle and proximal denti-setae slender, bifid and pectinate. Medially with one pectinate, spine-like seta and four or five long, simple setae. Maxillary palp ca. 1.1× length of galea-lacinia; 2-segmented; palp segment II 1.2× length of segment I; setae on maxillary palp fine, simple, scattered over surface of segments I and II; apex of last segment without excavation at inner distolateral margin, apically constricted.
Labium (Fig. 8h). Glossa basally broad, narrowing toward apex; shorter than paraglossa; inner margin with ten spine-like setae, increasing in length distally; apex with three long, robust, pectinate setae and one short, robust seta; outer margin with six or seven spine-like setae; ventral surface with fine, simple, scattered setae. Paraglossa sub-rectangular, curved inward; apex rounded; with three rows of long, robust, distally pectinate setae in apical area and two or three short, simple setae in anteromedial area; dorsally with a row of six or seven long, spine-like setae near inner margin. Labial palp with segment I 0.9× length of segments II and III combined. Segment I ventrally with short, fine, simple setae. Segment II with narrow, extended, distomedial protuberance; distomedial protuberance 0.6× width of base of segment III; ventral surface with short, simple setae; dorsally with three or four spine-like setae near outer margin. Segment III slightly pentagonal; length 0.9× width; ventrally covered with short, spine-like, simple setae and short, fine, simple setae.
Hind protoptera (Fig. 9i) absent.
Foreleg (Fig. 9a–d). Ratio of foreleg segments 1.3:1.0:0.6:0.2. Femur. Length ca. 4× maximum width. Dorsal margin with a row of 19–23 curved, spine-like, apically rounded setae; length of setae 0.1× maximum width of femur. Apex rounded, with a pair of spine-like setae and some short, stout setae. Many stout, lanceolate setae scattered along ventral margin; femoral patch present. On posterior side apically with stout setae. Tibia. Dorsal margin with a row of short, spine-like setae and fine, simple setae. Ventral margin with a row of short, curved, spine-like setae, on apex some longer setae and a tuft of fine, simple setae. Anterior surface scattered with stout, lanceolate setae. Patellotibial suture present on basal 1/2. Tarsus. Dorsal margin with a row of short, spine-like setae and fine, simple setae. Ventral margin with a row of curved, spine-like setae. Claw with one row of 10–12 denticles; distally pointed; with ca. six stripes; subapical setae absent.
Middle and hind legs (Fig. 9d). As foreleg, also with femoral patch. Stout setae on apex of posterior side present on middle leg and absent on hind leg.
Terga (Fig. 9e). Surface with irregular rows of U-shaped scale bases and scattered micropores. Posterior margin of tergum IV with triangular spines, longer than wide.
Gills (Fig. 9f). Present on segments II–VII. Margin with small denticles intercalating fine simple setae. Tracheae extending from main trunk to inner and outer margins. Gill IV as long as length of segments V and VI combined. Gill VII as long as length of segments VIII and 2/3 IX combined.
Paraproct (Fig. 9g). Distally slightly expanded, with 38–48 stout, marginal spines. Surface scattered with U-shaped scale bases and micropores. Cercotractor with numerous small, marginal spines.
Etymology
Dedicated to the indigenous Hattam people from West Papua.
Distribution
Indonesia: Papua Barat (Fig. 21c).
Biological aspects
The specimens were collected in a forest stream at an altitude of 500 m.
Labiobaetis werneri sp. nov.
Type material
Holotype. Papua New Guinea • larva; Gulf, Marawaka, Mala; 07°05'40"S, 145°44'28"E; 1400 m; 11.xi.2006; leg. Balke and Kinibel; (PNG 90); on slide; GenBank MW868307; GBIFCH00763699;
Diagnosis
Larva. Following combination of characters: A) dorsal surface of labrum with submarginal arc of one plus six or seven long, simple setae (Fig. 10a); B) labial palp segment II with rather narrow thumb-like, distomedial protuberance, segment III sub-rectangular (Fig. 10h); C) fore femur rather broad, length 2.7× maximum width, dorsal margin with 25–33 spine-like setae plus additional setae near margin (Fig. 11a); D) hind protoptera absent; E) six pairs of gills (gill I absent), anal margin with both short and long setae (Fig. 11e); F) paraproct distally not expanded, with 18–28 stout, marginal spines (Fig. 11f).
Description
Larva (Figs 10, 11, 19a, b). Body length 5.7–6.2 mm. Cerci broken, paracercus ca. 0.4× body length. Antenna: approx. 2.5× as long as head length.
Colouration (Fig. 19a, b). Head, thorax and abdomen dorsally brown, with pattern as in Fig. 19a. Head, thorax and abdomen ventrally light brown, abdominal sternites VI–IX darker, as in Fig. 19b. Legs light brown; femur with dorsomedial and apical brown spots; tarsus distally brown. Caudalii light brown.
Antenna (Fig. 11g) with scape and pedicel sub cylindrical, without distolateral process at scape.
Labrum (Fig. 10a). Sub-rectangular, length 0.7× maximum width. Distal margin with medial emargination and a small process. Dorsally with medium, fine, simple setae scattered over surface; submarginal arc of setae composed of one plus six or seven long, simple setae. Ventrally with marginal row of setae composed of anterolateral long, feathered setae and medial long, bifid, pectinate setae; ventral surface with ca. seven short, spine-like setae near lateral and anterolateral margin.
Right mandible (Fig. 10b, c). Incisor and kinetodontium fused. Incisor with four denticles; kinetodontium with three denticles, inner margin of innermost denticle with a row of thin setae. Prostheca robust, apically denticulate. Margin between prostheca and mola almost straight. Tuft of setae at apex of mola present.
Left mandible (Fig. 10d, e). Incisor and kinetodontium fused. Incisor with four denticles; kinetodontium with three denticles. Prostheca robust, apically with small denticles and comb-shaped structure. Margin between prostheca and mola straight. Subtriangular process long and slender, above level of area between prostheca and mola. Denticles of mola apically constricted. Tuft of setae at apex of mola present.
Both mandibles with lateral margins almost straight. Basal half with fine, simple setae scattered over dorsal surface.
Hypopharynx and superlinguae (Fig. 10f). Lingua approx. as long as superlinguae. Lingua longer than broad; distal half laterally slightly expanded; medial tuft of stout setae well developed and short. Superlinguae distally rounded; lateral margins rounded; fine, long, simple setae along distal margin.
Maxilla (Fig. 10g). Galea-lacinia ventrally with two simple, apical setae under canines. Inner dorsal row of setae with three denti-setae, distal denti-seta tooth-like, middle and proximal denti-setae slender, bifid and pectinate. Medially with one pectinate, spine-like seta and 7–9 long, simple setae. Maxillary palp ca. 1.3× length of galea-lacinia; 2-segmented; palp segment II 1.5× length of segment I; setae on maxillary palp fine, simple, scattered over surface of segments I and II; apex of last segment without excavation at inner distolateral margin, apically constricted.
Labium (Fig. 10h). Glossa basally broad, narrowing toward apex; shorter than paraglossa; inner margin with nine spine-like setae, increasing in length distally; apex with three long, robust, pectinate setae and one short, robust seta; outer margin with seven or eight spine-like setae; ventral surface with fine, simple, scattered setae. Paraglossa sub-rectangular, curved inward; apex rounded; with three rows of long, robust, distally pectinate setae in apical area, three or four short, simple setae in anteromedial area and one short, simple seta in posterolateral area; dorsally with a row of five long, spine-like setae near inner margin. Labial palp with segment I 1.1× length of segments II and III combined. Segment I ventrally with short, fine, simple setae. Segment II with narrow thumb-like, distomedial protuberance; distomedial protuberance 0.5× width of base of segment III; ventral surface with short, simple setae; dorsally with 3–5 spine-like setae near outer margin. Segment III sub-rectangular; length 1.1× width; ventrally covered with short, spine-like, simple setae and short, fine, simple setae.
Hind protoptera (Fig. 11h) absent.
Foreleg (Fig. 11a–c). Ratio of foreleg segments 1.6:1.0:0.9:0.3. Femur. Length 2.7× maximum width. Dorsal margin with a row of 25–33 curved, spine-like setae and additional setae near margin; length of setae 0.16× maximum width of femur. Apex rounded, with a pair of spine-like setae and some short, stout setae. Many stout, lanceolate setae scattered along ventral margin and some scattered on surface; femoral patch present. On posterior side apically with stout setae. Tibia. Dorsal margin with a row of short to medium, spine-like setae. Ventral margin with a row of short, curved, spine-like setae, on apex some longer setae and a tuft of fine, simple setae. Anterior surface scattered with stout, lanceolate setae. Patellotibial suture present on basal 1/2. Tarsus. Dorsal margin with a row of short, spine-like setae and fine, simple setae. Ventral margin with a row of curved, spine-like setae. Claw with one row of 9–12 denticles; distally pointed; with ca. five stripes; subapical setae absent.
Middle and hind legs (Fig. 11c). As foreleg, also with femoral patch. Stout setae on apex of posterior side present on middle leg and absent on hind leg.
Terga (Fig. 11d). Surface with irregular rows of U-shaped scale bases and scattered micropores. Posterior margin of tergum IV with triangular spines, wider than long.
Gills (Fig. 11e). Present on segments II–VII. Costal margin with small denticles intercalating short, fine simple setae; anal margin with small denticles, intercalating both short and long, fine, simple setae. Tracheae extending from main trunk to inner and outer margins. Gill IV as long as length of segments V and 2/3 VI combined. Gill VII as long as length of segments VIII and 1/2 IX combined.
Paraproct (Fig. 11f). Distally not expanded, with 18–28 stout, marginal spines. Surface scattered with U-shaped scale bases and fine, simple setae. Cercotractor with numerous small, marginal spines.
Etymology
Dedicated to Werner Horzel, the late stepfather of the first author.
Distribution
Papua New Guinea (Fig. 21c).
Biological aspects
The specimens were collected at altitudes from 1150 m to 1800 m.
Labiobaetis dendrisetis group of species (Kaltenbach et al. 2020 )
The dendrisetis group can be recognised by the following combination of characters: A) dorsal surface of labrum with submarginl arc of dendritic setae; B) labial palp segment II with narrow thumb-like protuberance; C) labial palp segment III broad, rounded; D) seven pairs of gills.
The L. dendrisetis group is present in New Guinea and the Philippines; it includes the following species:
Labiobaetis dalisay Kaltenbach, Garces & Gattolliat, 2020
Labiobaetis dendrisetis Kaltenbach & Gattolliat, 2018
Labiobaetis dendrisetis
Material examined
Papua New Guinea • 9 larvae; Central Prov., Kokoda Trek; 09°00'20"S, 147°44'15"E; 1390 m; i.2008; leg. Posman; (PNG 173); 2 on slides; GenBank MW868310; GBIFCH00763706, 592769;
Diagnosis
Larva. Following combination of characters: A) dorsal surface of labrum with submarginal arc of long, dendritic setae setae (
Due to the limited material in the type series (holotype and one paratype), a few parts of the original description were missing, incomplete or have to be corrected:
Complementary description
Larva (Figs 12, 19c, d). Body length 4.1–5.3 mm. Cerci broken, paracercus ca. half body length.
Colouration (Fig. 19c, d). Head dorsally ochreous, thorax and abdomen dorsally brown, with pattern as in Fig. 19c. Head, thorax and abdomen ventrally ecru, abdominal sternites VI–VIII dark brown (Fig. 19d). Legs light brown, femur with distomedial brown streak. Caudalii light brown.
Maxillary palp (Fig. 12a) ca. 1.1× length of galea-lacinia; 2-segmented; palp segment II approx. as long as segment I; apex of last segment with slight excavation at inner distolateral margin, apically rounded.
Gills (Fig. 12b). Present on segments I–VII. Margin with small denticles intercalating fine simple setae. Tracheae extending from main trunk to inner and outer margins. Gill I as long as length of segment II. Gill IV as long as length of segments V, VI and 2/3 VII combined. Gill VII as long as length of segments VIII, IX and 1/3 X combined.
Distribution
Papua New Guinea (Fig. 21c).
Labiobaetis seramensis group of species (Kaltenbach and Gattolliat 2019 )
The seramensis group is recognised by the following combination of characters: A) dorsal surface of labrum with submarginal arc of simple setae; B) labial palp segment II with narrow or rather narrow, thumb-like distomedial protuberance, segment III broad, rounded; C) femur dorsally with dense setation; middle and hind leg with reduced femoral patch; D) six pairs of gills (gill I absent); E) hind protoptera absent; F) distolateral process at scape absent.
The L. seramensis group is reported from Seram (Indonesia) and New Guinea (Indonesia: Papua Barat), it includes the following species:
Labiobaetis arfak sp. nov. (New Guinea)
Labiobaetis onim sp. nov. (New Guinea)
Labiobaetis seramensis Kaltenbach & Gattolliat, 2019 (Seram)
Labiobaetis wahai Kaltenbach & Gattolliat, 2019 (Seram)
Key to the species of the Labiobaetis seramensis group (larvae)
| 1 | Dorsal margin of femur with row of more than 70 long, curved, spine-like setae (Fig. 16a); posterior margin of tergite IV with discontinued row of triangular spines (Fig. 16e) | L. onim sp. nov. |
| – | Dorsal margin of femur with less than 25 curved, spine-like setae; posterior margin of tergite IV with continued row of triangular spines (Fig. 14e) | 2 |
| 2 | Abdominal tergites dark brown, segments V and VI yellow brown (Fig. 20a); posterolateral margins of tergites VIII and IX with two long, pointed spines (Fig. 14f) | L. arfak sp. nov. |
| – | Abdominal tergites light brown ( |
3 |
| 3 | Posterior margins of tergites with triangular spines, wider than long ( |
L. seramensis |
| – | Posterior margins of tergites with triangular spines, longer than wide ( |
L. wahai |
Labiobaetis arfak sp. nov.
Type material
Holotype. Indonesia • larva; Papua Barat, River Je, Loc. Arfak, East of Amber village; 01°10'59"S, 133°56'51"E; 1200 m; 16.vi.2016, leg. Sumoked; on slide; GBIFCH00592770;
Diagnosis
Larva. Following combination of characters: A) dorsal surface of labrum with submarginal arc of one plus two long, simple setae (Fig. 13a); B) labial palp segment II with narrow thumb-like, distomedial protuberance, segment III broad, rounded (Fig. 13h); C) fore femur rather broad, length 2.7× maximum width, dorsal margin with 17–20 spine-like setae plus a second row of spine-like setae near margin (Fig. 14a); D) hind protoptera absent; E) six pairs of gills (gill I absent); F) tergites VIII and IX posterolaterally with two long spines (Fig. 14f); G) paraproct distally not expanded, with ca. eight stout, marginal spines (Fig. 14h).
Description
Larva (Figs 13, 14, 20a, b). Body length 3.3–4.4 mm. Cerci ca. 2/3 of body length, paracercus ca. 2/3 of cerci length. Antenna: approx. 2.5× as long as head length.
Colouration (Fig. 20a, b). Head, thorax and abdomen dorsally dark brown, abdominal tergites V and VI yellow brown (Fig. 20a). Thorax ventrally ecru, abdomen ventrally brown, with pattern as in Fig. 20b. Legs light brown; femur dorsally and ventrally dark brown, basally and distomedially with dark brown areas (Fig. 20b). Caudalii light brown, basally brown.
Antenna (Fig. 14i) with scape and pedicel sub cylindrical, without distolateral process at scape.
Labrum (Fig. 13a). Sub-rectangular, length 0.6× maximum width. Distal margin with medial emargination and a small process. Dorsally with medium, fine, simple setae scattered over surface; submarginal arc of setae composed of one plus two long, simple setae. Ventrally with marginal row of setae composed of anterolateral long, feathered setae and medial long, bifid, pectinate setae; ventral surface with ca. three short, spine-like setae near lateral and anterolateral margin.
Right mandible (Fig. 13b, c). Incisor and kinetodontium fused. Incisor with five denticles; kinetodontium with three denticles, inner margin of innermost denticle with a row of thin setae. Prostheca robust, apically denticulate. Margin between prostheca and mola convex, with minute denticles. Tuft of setae at apex of mola present.
Left mandible (Fig. 13d, e). Incisor and kinetodontium fused. Incisor with five denticles; kinetodontium with three denticles. Prostheca robust, apically with small denticles and comb-shaped structure. Margin between prostheca and mola almost straight, with few minute denticles. Subtriangular process long and slender, above level of area between prostheca and mola. Denticles of mola apically constricted. Tuft of setae at apex of mola absent.
Both mandibles with lateral margins almost straight. Basal half with fine, simple setae scattered over dorsal surface.
Hypopharynx and superlinguae (Fig. 13f). Lingua shorter than superlingua. Lingua longer than broad; distal half laterally slightly expanded; medial tuft of stout setae well developed and long. Superlinguae distally rounded; lateral margins rounded; fine, long, simple setae along distal margin.
Maxilla (Fig. 13g). Galea-lacinia ventrally with two simple, apical setae under canines. Inner dorsal row of setae with three denti-setae, distal denti-seta tooth-like, middle and proximal denti-setae slender, bifid and pectinate. Medially with one spine-like seta and four long, simple setae. Maxillary palp ca. 1.3× length of galea-lacinia; 2-segmented; palp segment II 1.5× length of segment I; setae on maxillary palp fine, simple, scattered over surface of segment II; apex of last segment without excavation at inner distolateral margin, apically rounded.
Labium (Fig. 13h). Glossa basally broad, narrowing toward apex; shorter than paraglossa; inner margin with 3–5 spine-like setae, distalmost seta much longer; apex with two long and one medium robust, pectinate setae; outer margin with three or four spine-like setae; ventral surface with fine, simple, scattered setae. Paraglossa sub-rectangular, curved inward; apex rounded; with three rows of long, robust, distally pectinate setae in apical area, one or two short, simple setae in anteromedial area and one short, simple seta in posterolateral area; dorsally with a row of four long, spine-like setae near inner margin. Labial palp with segment I 0.9× length of segments II and III combined. Segment I ventrally with short, fine, simple setae. Segment II with narrow thumb-like, distomedial protuberance; distomedial protuberance 0.4× width of base of segment III; ventral surface with short, simple setae; dorsally with two or three spine-like setae near outer margin. Segment III broad, rounded; length 0.9× width; ventrally covered with short, spine-like, simple setae and short, fine, simple setae.
Hind protoptera (Fig. 14j) absent.
Foreleg (Fig. 14a, b). Ratio of foreleg segments 1.3:1.0:0.5:0.2. Femur. Length 2.7× maximum width. Dorsal margin with a row of 17–20 curved, spine-like setae and a second row of spine-like setae near margin; length of setae 0.25× maximum width of femur. Apex rounded, with one or two pairs of spine-like setae and some short, stout setae. Many stout, lanceolate setae scattered along ventral margin; femoral patch absent. Tibia. Dorsal margin with a row of short, spine-like setae and fine, simple setae. Ventral margin with a row of short, curved, spine-like setae. Anterior surface scattered with few stout, lanceolate setae. Patellotibial suture present on basal 1/2 area. Tarsus. Dorsal margin with a row of short, spine-like setae. Ventral margin with a row of curved, spine-like setae. Claw with one row of 9–11 denticles; distally pointed; with ca. five stripes; subapical setae absent.
Middle and hind legs (Fig. 14c, d). As foreleg, but with reduced femoral patch.
Terga (Fig. 14e, f). Surface with irregular rows of U-shaped scale bases and scattered micropores. Posterior margin of tergum IV with triangular spines, wider than long. Posterolateral margins of terga VIII and IX with two long, pointed spines.
Gills (Fig. 14g). Present on segments II–VII. Margin with small denticles intercalating fine simple setae. Tracheae extending from main trunk to inner and outer margins. Gill IV as long as length of segments V and 2/3 VI combined. Gill VII as long as length of segments VIII and 2/3 IX combined.
Paraproct (Fig. 14h). Distally not expanded, with ca. eight stout, marginal spines. Surface scattered with U-shaped scale bases, micropores and fine, simple setae. Cercotractor with numerous small, marginal spines.
Etymology
Dedicated to the indigenous Arfak people of Papua Barat, where the type locality is located.
Distribution
Indonesia: Papua Barat (Fig. 21c).
Biological aspects
The specimens were collected at an altitude of 1200 m, together with L. onim sp. nov.
Labiobaetis onim sp. nov.
Type material
Holotype. Indonesia • larva; Papua Barat, River Je, Loc. Arfak, East of Amber village; 01°10'59"S, 133°56'51"E; 1200 m; 16.vi.2016, leg. Sumoked; on slide; GBIFCH00763713;
Diagnosis
Larva. Following combination of characters: A) dorsal surface of labrum with submarginal arc of one plus two long, simple setae (Fig. 15a); B) labial palp segment II with short thumb-like (atypical for the group), distomedial protuberance, segment III broad, rounded (Fig. 15h); C) fore femur rather broad, length ca. 3× maximum width, dorsal margin with more than 70 long, curved spine-like setae plus some additional spine-like setae near margin (Fig. 16a); D) hind protoptera absent; E) six pairs of gills (gill I absent); F) paraproct distally not expanded, with 8–12 stout, marginal spines (Fig. 16g).
Description
Larva (Figs 15, 16, 20c, d). Body length 5.6–5.9 mm. Cerci ca. 1/2 of body length, paracercus ca. 2/3 of cerci length. Antenna: approx. twice as long as head length.
Colouration (Fig. 20c, d). Head dorsally light brown, thorax and abdomen dorsally dark brown, with light brown pattern on thorax as in Fig. 20c, abdominal segment I light brown and abdominal segments V–VII orange. Head, thorax, and abdomen ventrally light brown, with pattern as in fig. 20d, abdominal segments V–VII light orange and abdominal segments VIII–X dark brown. Legs ecru, caudalii ecru.
Antenna (Fig. 16h) with scape and pedicel sub cylindrical, without distolateral process at scape.
Labrum (Fig. 15a). Sub-rectangular, length 0.6× maximum width. Distal margin with medial emargination and a small process. Dorsally with medium, fine, simple setae scattered over surface; submarginal arc of setae composed of one plus two long, simple setae; large distance between both arc setae. Ventrally with marginal row of setae composed of anterolateral long, feathered setae and medial long, bifid, pectinate setae; ventral surface with ca. five short, spine-like setae near lateral and anterolateral margin.
Right mandible (Fig. 15b, c). Incisor and kinetodontium fused. Incisor with six denticles; kinetodontium with three denticles, inner margin of innermost denticle with a row of thin setae. Prostheca robust, apically denticulate. Margin between prostheca and mola slightly convex. Tuft of setae at apex of mola present.
Left mandible (Fig. 15d, e). Incisor and kinetodontium fused. Incisor with five denticles; kinetodontium with three denticles. Prostheca robust, apically with small denticles and comb-shaped structure. Margin between prostheca and mola straight. Subtriangular process long and slender, above level of area between prostheca and mola. Denticles of mola apically constricted. Tuft of setae at apex of mola absent.
Both mandibles with lateral margins almost straight. Basal half with fine, simple setae scattered over dorsal surface.
Hypopharynx and superlinguae (Fig. 15f). Lingua shorter than superlingua. Lingua longer than broad; distal half laterally slightly expanded; medial tuft of stout setae well developed and long. Superlinguae distally rounded; lateral margins rounded; fine, long, simple setae along distal margin.
Maxilla (Fig. 15g). Galea-lacinia ventrally with two simple, apical setae under canines. Inner dorsal row of setae with three denti-setae, distal denti-seta tooth-like, middle and proximal denti-setae slender, bifid and pectinate. Medially with one pectinate, spine-like seta and three long, simple setae. Maxillary palp ca. 1.3× length of galea-lacinia; 2-segmented; palp segment II 1.1× length of segment I; setae on maxillary palp fine, simple, scattered over surface of segments I and II; apex of last segment without excavation at inner distolateral margin, apically rounded.
Labium (Fig. 15h). Glossa basally broad, narrowing toward apex; shorter than paraglossa; inner margin with one long, spine-like seta; apex with two long and one medium robust, pectinate setae; outer margin with five or six spine-like setae; ventral surface with fine, simple, scattered setae. Paraglossa sub-rectangular, curved inward; apex rounded; with three rows of long, robust, distally pectinate setae in apical area, sometimes one short, simple seta in anteromedial area, and one short, simple seta in posteromedial area; dorsally with a row of three long, spine-like setae near inner margin. Labial palp with segment I approx. as long as segments II and III combined. Segment I ventrally with short, fine, simple setae. Segment II with short thumb-like, distomedial protuberance; distomedial protuberance 0.3× width of base of segment III; ventral surface with short, simple setae; dorsally with one spine-like seta near outer margin. Segment III broad, rounded; length 0.7× width; ventrally covered with short, spine-like, simple setae and short, fine, simple setae.
Hind protoptera absent.
Foreleg (Fig. 16a, b). Ratio of foreleg segments 1.2:1.0:0.5:0.2. Femur. Length ca. 3× maximum width. Dorsal margin with a dense row of more than 70 long, curved, spine-like setae and distally some additional long, spine-like setae near margin; length of setae 0.40× maximum width of femur. Apex rounded, with some short, stout setae. Many stout, lanceolate setae scattered along ventral margin; femoral patch absent. Tibia. Dorsal margin with a dense row of long, spine-like setae. Ventral margin with a row of short, curved, spine-like setae. Anterior surface scattered with short, stout, lanceolate setae. Patellotibial suture present on basal 1/2 area. Tarsus. Dorsal margin with a row of short, spine-like setae. Ventral margin with a row of short, curved, spine-like setae, distalmost seta much longer. Claw with one row of ten or eleven denticles; distally pointed; with two or three stripes; subapical setae absent.
Middle and hind legs (Fig. 16c, d). As foreleg, but with reduced femoral patch.
Terga (Fig. 16e). Surface with scattered scales and micropores. Posterior margin of tergum IV with discontinuous row of triangular spines, spines wider than long. Triangular spines present on segments IV–VII, absent on segments I–III.
Gills (Fig. 16f). Present on segments II–VII. Margin with small denticles intercalating fine simple setae. Tracheae partly extending from main trunk to inner and outer margins. Gill IV as long as length of segments V and 1/2 VI combined. Gill VII slightly longer than length of segment VIII.
Paraproct (Fig. 16g). Distally not expanded, with 8–12 stout, marginal spines, partly with split tips. Surface scattered with scales and micropores. Cercotractor with numerous small, marginal spines, partly with split tips.
Etymology
Dedicated to the indigenous Onim people of Papua Barat, where the type locality is located.
Distribution
Indonesia: Papua Barat (Fig. 21c).
Biological aspects
The specimens were collected at an altitude of 1200 m, together with L. arfak sp. nov.
Discussion
Assignment to Labiobaetis
For the assignment of the new species to Labiobaetis we refer to
Labiobaetis catadupa sp. nov. group
This group is formed for two new species sharing distally very slender glossae, much shorter than paraglossae; an extended thumb-like, hooked protuberance of labial palp segment II; the presence of a long subapical seta on the claw and opposite a rudimentary subapical seta; and the absence of the first pair of gills, hind protoptera and a distolateral process at scape (Figs 1i, 2f, g, 4, 6h, 7h, i). Also, both have a submarginal arc of setae dorsally on the labrum composed of feathered setae. However, the type of these setae of L. toraja sp. nov. (Fig. 6a) is common in Labiobaetis, present in all species of the groups operosus and difficilis from Southeast Asia and in almost all species of the Afrotropical realm (
The genetic distance (COI, Kimura 2-parameter) between L. catadupa sp. nov. and L. toraja sp. nov. is 23% and thus well in line with distances found between different species of Labiobaetis in Southeast Asia (15%–27% in the Philippines,
Labiobaetis claudiae group
Based on the discovery of two further species of this group, L. hattam sp. nov. and L. werneri sp. nov., we adapted the diagnosis of the group (see in the results section) and added another species, L. centralensis, already described earlier (
| Species | Locality | GPS coordinates |
|---|---|---|
| L. catadupa sp. nov. | Brunei: Temburong National Park | 04°32'49"N, 115°09'30"E |
| 04°33'10"N, 115°09'20"E | ||
| 04°32'42"N, 115°09'31"E | ||
| 04°32'23"N, 115°09'34"E | ||
| 04°32'51"N, 115°09'25"E | ||
| 04°32'56"N, 115°09'27"E | ||
| 04°33'39"N, 115°08'51"E | ||
| 04°33'39"N, 115°08'54"E | ||
| L. toraja sp. nov. | Sulawesi | 01°19'35"S, 120°18'40"E |
| L. hattam sp. nov. | Papua Barat | 00°52'29"S, 132°46'06"E |
| L. werneri sp. nov. | Papua New Guinea | 07°01'42"S, 145°49'48"E |
| Papua New Guinea | 07°05'40"S, 145°44'28"E | |
| Papua New Guinea | 06°51'04"S, 146°48'04"E | |
| Papua New Guinea | 07°51'02"S, 147°07'00"E | |
| L. dendrisetis | Papua New Guinea: Simbu Prov. | 05°49' 00"S, 145°04'30"E |
| Papua New Guinea: Central Prov. | 08°31'35"S, 147°14'06"E | |
| Papua New Guinea: Central Prov. | 08°31'17"S, 147°13'41"E | |
| Papua New Guinea: Central Prov. | 09°00'20"S, 147°44'15"E | |
| L. arfak sp. nov. | Papua Barat | 01°10'59"S, 133°56'51"E |
| L. onim sp. nov. | Papua Barat | 01°10'59"S, 133°56'51"E |
The interspecific genetic distances between the species of the claudiae group are rather high, between 18% and 27% (Table 3), which is in line with the values reported for other Labiobaetis species in New Guinea (13%–28%;
Intraspecific (bold) and interspecific genetic distances of the species of the L. claudiae group (COI; Kimura 2-parameter).
| 1 | 2 | 3 | 4 | 5 | 6 | 7 | 8 | 9 | ||
|---|---|---|---|---|---|---|---|---|---|---|
| 1 | L. academicus | |||||||||
| 2 | L. academicus | 0.00 | ||||||||
| 3 | L. centralensis | 0.22 | 0.22 | |||||||
| 4 | L. centralensis | 0.23 | 0.23 | 0.01 | ||||||
| 5 | L. claudiae | 0.21 | 0.21 | 0.24 | 0.24 | |||||
| 6 | L. hattam sp. nov. | 0.23 | 0.23 | 0.23 | 0.23 | 0.23 | ||||
| 7 | L. stagnum | 0.20 | 0.20 | 0.26 | 0.27 | 0.23 | 0.23 | |||
| 8 | L. werneri sp. nov. | 0.19 | 0.19 | 0.25 | 0.25 | 0.20 | 0.21 | 0.23 | ||
| 9 | L. werneri sp. nov. | 0.18 | 0.18 | 0.25 | 0.25 | 0.20 | 0.21 | 0.22 | 0.03 | |
| 10 | L. werneri sp. nov. | 0.19 | 0.19 | 0.25 | 0.25 | 0.20 | 0.21 | 0.22 | 0.05 | 0.06 |
Labiobaetis seramensis group
Due to the discovery of two new species of the L. seramensis group in New Guinea, L. arfak sp. nov. and L. onim sp. nov., and the re-examination of the types of L. seramensis and L. wahai from Seram (Indonesia), we could complement the diagnosis of this group (see in section results). The presence of a reduced femoral patch on middle and hind legs and its absence on forelegs is identified as an additional character of the group. As femoral patches are generally rare in Labiobaetis in Southeast Asia and New Guinea, it is considered to be a strong character.
Labiobaetis onim sp. nov. has three remarkable characters, which are atypical for Labiobaetis: the femur has a very dense and long setation with more than 70 setae at the dorsal margin, the tibia also has a dense and rather long setation along its dorsal margin and the posterior margins of the tergites have a discontinuous row of triangular spines, similar as in Baetis noa Yanai & Gattolliat, 2018 from Israel (
Distribution of Labiobaetis in the Wallacea region
The L. seramensis group is present with two species on the island Seram (Indonesia, Moluccas) and with two others in New Guinea. Moreover, L. arfak sp. nov. from New Guinea is morphologically very similar to L. seramensis from Seram (
The Wallace Line is marking the eastern boundary of the Oriental fauna, and Lydekker’s Line is considered to be the western boundary of the strictly Australian fauna. The mixed zone in between is referred to as Wallacea by many biogeographers (
Independent from the situation in the Wallacea, there could have been a limited stepping stone exchange between the Philippines and New Guinea, as we found members of the groups vallus and dendrisetis in both these archipelagos (
Taking into account the extreme diversity in Southeast Asia and New Guinea, the rather poor collection activities in the past, with many still unexplored regions, and the obvious richness of Labiobaetis in this region, we have to expect many more species with further collections in the future.
Acknowledgements
We sincerely thank Kate Baker (University of Exeter, UK) for the collection of this precious material and to make it available to the Museum of Zoology Lausanne (
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