Research Article |
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Corresponding author: Chanaporn Suttinun ( chanaporn.s@cmu.ac.th ) Academic editor: Ben Price
© 2025 Thomas Kaltenbach, Jean-Luc Gattolliat, Boonsatien Boonsoong, Chanaporn Suttinun.
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, Boonsoong B, Suttinun C (2025) New species of Labiobaetis Novikova & Kluge from Thailand (Ephemeroptera, Baetidae). ZooKeys 1258: 213-276. https://doi.org/10.3897/zookeys.1258.166681
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Investigations of material collected by one of us (CS) between May 2017 and November 2023 from 70 localities in Thailand further increases our knowledge of the diversity of the genus Labiobaetis Novikova & Kluge, 1987 in Thailand and Southeast Asia in general. Eleven species have been identified using a combination of morphological and molecular analysis (COI). Nine are new to science, they are described and illustrated based on their larvae, and in one case, complemented by the male imago. Two of the new species belong to the L. batakorum species group, L. mon sp. nov., L. lahu sp. nov.; three to the L. numeratus species group, L. tenasserimensis sp. nov., L. angularis sp. nov., L. tonsator sp. nov.; one to the L. operosus species group, L. nisaratae sp. nov., and three to the L. sumigarensis species group, L. karen sp. nov., L. septem sp. nov., L. ranongensis sp. nov. A key to all species of Labiobaetis from continental Southeast Asia is provided. Additionally, the genetic distances (COI; Kimura-2 parameter) including all species treated in this study are discussed. The total number of Labiobaetis species worldwide is augmented to approximately 170.
COI, integrative taxonomy, mayflies, morphology, Southeast Asia
Southeast Asia is one of the regions with the highest biodiversity worldwide in general, and also specifically for mayflies. Considerable effort has been made in recent years to gain a better understanding of this yet understudied fauna, including studies focusing on the lesser known, but highly diverse mayfly family Baetidae. Particular emphasis was placed especially on Thailand, but also on the archipelagos of Indonesia and the Philippines, where new genera of Baetidae and many new species were discovered (e.g.
The genus Labiobaetis Novikova & Kluge, 1987 belongs to Baetidae, the most diverse family of Ephemeroptera. More than one fourth of all mayfly species worldwide (> 1200 species) are part of Baetidae (
Recently, a comprehensive study on the Baetidae of Thailand was conducted by
Given the extraordinary diversity in Thailand and despite the strong effort already done so far, it is reasonable to anticipate the discovery of numerous additional species through future studies in Thailand and in surrounding countries.
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.
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 v. 5.3 (http://www.heliconsoft.com). Photographs of larval parts on slides were taken with an Olympus BX43 microscope equipped with an Olympus SC 50 camera and the program Olympus CellSense v. 4.1. All photographs were subsequently enhanced with Adobe Photoshop Elements 13.
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
| Species | Specimen voucher | Specimen voucher | GPS coordinates | GenBank | GenSeq |
|---|---|---|---|---|---|
| catalogue # | catalogue # (slide) | # (COI) | Nomenclature | ||
| L. multus (Sumatra) | GBIFCH00235847 | – | 00°34'25"S, 100°43'54"E | MN167323 | genseq-4 COI |
| L. multus | GBIFCH00829311 | GBIFCH01556161 | 07°34'32"N, 99°47'13"E | PX067793 | genseq-4 COI |
| GBIFCH00980855 | – | 07°29'33"N, 99°46'26"E | PX067795 | genseq-4 COI | |
| GBIFCH00980853 | – | 07°43'44"N, 99°44'21"E | PX067794 | genseq-4 COI | |
| L. mon sp. nov. | GBIFCH00829315 | GBIFCH01556160 | 12°03'50"N, 99°37'39"E | PX067748 | genseq-2 COI |
| GBIFCH00829307 | GBIFCH00592518 | 13°30'33"N, 99°17'12"E | PX067749 | genseq-2 COI | |
| GBIFCH00980852 | – | 13°30'57"N, 99°20'40"E | PX067746 | genseq-2 COI | |
| GBIFCH00980854 | – | 12°58'42"N, 99°34'55"E | PX067745 | genseq-2 COI | |
| GBIFCH00980857 | – | 12°58'42"N, 99°34'55"E | PX067747 | genseq-1 COI | |
| GBIFCH00829296 | – | 12°52'36"N, 102°05'48"E | PX067744 | genseq-2 COI | |
| L. lahu sp. nov. | GBIFCH00829303 | GBIFCH01223071 | 16°48'16"N, 99°01'19"E | PX067750 | genseq-2 COI |
| GBIFCH00829301 | GBIFCH01223072 | 16°46'53"N, 99°01'16"E | PX067753 | genseq-2 COI | |
| GBIFCH00829318 | GBIFCH01223073 | 16°46'53"N, 99°01'16"E | PX067758 | genseq-2 COI | |
| GBIFCH00829302 | GBIFCH00596154 | 16°46'53"N, 99°01'16"E | PX067754 | genseq-1 COI | |
| GBIFCH00829298 | GBIFCH00607175 | 17°18'04"N, 101°46'33"E | PX067752 | genseq-2 COI | |
| GBIFCH00829306 | GBIFCH00607176 | 19°17'02"N, 98°58'05"E | PX067755 | genseq-2 COI | |
| GBIFCH00829295 | – | 16°42'04"N, 98°30'41"E | PX067751 | genseq-2 COI | |
| GBIFCH00980856 | – | 19°17'02"N, 98°58'05"E | PX067757 | genseq-2 COI | |
| GBIFCH00980858 | – | 19°17'02"N, 98°58'05"E | PX067759 | genseq-2 COI | |
| GBIFCH00980851 | – | 13°30'57"N, 99°20'40"E | PX067756 | genseq-2 COI | |
| L. tenasserimensis sp. nov. | GBIFCH00829286 | – | 13°31'27"N, 99°14'39"E | PX067774 | genseq-2 COI |
| GBIFCH00829275 | – | 13°31'27"N, 99°14'39"E | PX067772 | genseq-2 COI | |
| GBIFCH00829285 | – | 09°41'27"N, 98°35'19"E | PX067773 | genseq-2 COI | |
| L. angularis sp. nov. | GBIFCH00829290 | – | 12°58'42"N, 99°34'55"E | PX067788 | genseq-2 COI |
| GBIFCH00829278 | – | 13°24'22"N, 99°16'44"E | PX067779 | genseq-2 COI | |
| GBIFCH00829279 | – | 12°58'42"N, 99°34'55"E | PX067780 | genseq-2 COI | |
| GBIFCH00829287 | – | 12°03'50"N, 99°37'39"E | PX067786 | genseq-2 COI | |
| GBIFCH00829277 | – | 12°38'14"N, 99°30'59"E | PX067777 | genseq-2 COI | |
| GBIFCH00829280 | – | 12°46'03"N, 99°34'54"E | PX067781 | genseq-2 COI | |
| GBIFCH00829283 | – | 13°30'57"N, 99°20'40"E | PX067784 | genseq-2 COI | |
| GBIFCH00829274 | – | 12°46'03"N, 99°34'54"E | PX067778 | genseq-2 COI | |
| GBIFCH00829284 | – | 14°34'58"N, 98°34'52"E | PX067785 | genseq-2 COI | |
| GBIFCH00829273 | – | 09°42'22"N, 98°34'39"E | PX067775 | genseq-2 COI | |
| GBIFCH00829282 | – | 13°30'33"N, 99°17'12"E | PX067783 | genseq-2 COI | |
| GBIFCH00829289 | – | 13°28'41"N, 99°14'55"E | PX067787 | genseq-2 COI | |
| GBIFCH00829291 | – | 13°24'22"N, 99°16'44"E | PX067789 | genseq-2 COI | |
| GBIFCH00829281 | – | 17°16'11"N, 101°35'51"E | PX067782 | genseq-2 COI | |
| GBIFCH00829276 | – | 12°52'36"N, 102°05'48"E | PX067776 | genseq-2 COI | |
| L. tonsator sp. nov. | GBIFCH00829272 | – | 07°11'43"N, 100°04'18"E | PX067791 | genseq-1 COI |
| L. cf. paraoperosus | GBIFCH00829310 | GBIFCH01223074 | 10°03'54"N, 98°40'13"E | PX067797 | genseq-4 COI |
| GBIFCH00829312 | GBIFCH00596156 | 10°03'54"N, 98°40'13"E | PX067796 | genseq-4 COI | |
| L. nisaratae sp. nov. | GBIFCH00829316 | GBIFCH01223084 | 14°34'58"N, 98°34'52"E | PX067790 | genseq-2 COI |
| L. karen sp. nov. | GBIFCH00829300 | GBIFCH00592515 | 16°46'53"N, 99°01'16"E | PX067760 | genseq-2 COI |
| GBIFCH00980859 | – | 14°34'58"N, 98°34'52"E | PX067761 | genseq-2 COI | |
| L. septem sp. nov. | GBIFCH00829297 | GBIFCH00596155 | 14°34'58"N, 98°34'52"E | PX067765 | genseq-2 COI |
| GBIFCH00829292 | GBIFCH01221815 | 16°46'53"N, 99°01'16"E | PX067762 | genseq-2 COI | |
| GBIFCH00829294 | GBIFCH01221816 | 17°16'11"N, 101°35'51"E | PX067763 | genseq-2 COI | |
| GBIFCH00975856 | – | 16°46'53"N, 99°01'16"E | PX067766 | genseq-4 COI | |
| GBIFCH00975857 | – | 16°46'53"N, 99°01'16"E | PX067767 | genseq-4 COI | |
| GBIFCH00975858 | – | 16°46'53"N, 99°01'16"E | PX067768 | genseq-4 COI | |
| GBIFCH00980863 | GBIFCH01221817 | 16°44'28"N, 99°02'43"E | PX067764 | genseq-4 COI | |
| L. cf. septem sp. nov. | GBIFCH00975853 | – | 19°19'19"N, 98°52'51"E | PX067798 | genseq-4 COI |
| GBIFCH00975854 | – | 19°19'19"N, 98°52'51"E | PX067799 | genseq-4 COI | |
| L. ranongensis sp. nov. | GBIFCH00980860 | – | 09°52'08"N, 98°37'32"E | PX067769 | genseq-2 COI |
| GBIFCH00980864 | – | 09°43'25"N, 98°36'29"E | PX067770 | genseq-1 COI | |
| GBIFCH00975861 | – | 09°42'22"N, 98°34'39"E | PX067771 | genseq-2 COI |
The distribution maps were generated with the program SimpleMappr (https://simplemappr.net,
The dichotomous key was elaborated with the support of the program DKey v. 1.3.0 (http://drawwing.org/dkey,
The terminology follows
batakorum group
numeratus group
operosus group
sumigarensis group
Diagnosis. Dorsal surface of labrum with submarginal arc of simple setae, 1st and 2nd setae after submedian seta standing closely together; labial palp segment II with large, lobed distomedial protuberance; left mandible without setae at apex of mola; lingua of hypopharynx with well developed, short medial tuft of setae; distolateral process at scape well developed; paraproct distally not expanded; hind protoptera absent (L. batakorum species group s. str.) or present (L. multus and closely related species L. mon sp. nov. and L. lahu sp. nov.).
Probably, the L. multus species group should be included into the L. tricolor species group (
The following species are known from continental Southeast Asia: Labiobaetis multus (Müller-Liebenau, 1984) (new assignment to the group), Labiobaetis mon sp. nov., Labiobaetis lahu sp. nov.
Baetis multus Müller-Liebenau, 1984: 263.
Labiobaetis multus
(Müller-Liebenau):
Thailand • 1 larva; Trang Prov., Na Yong Distr., Wang Nam Rab resort; 07°34'32"N, 99°47'13"E; 142 m; 18.v.2017; leg. C. Suttinun; on slide; GBIFCH01556161;
Larva. Following combination of characters differentiate L. multus from other species of the group batakorum: head and thorax dorsally beige with grey-brown markings, abdomen dorsally grey-brown, laterally whitish with black markings, terga V and X brighter (Fig.
The species differs from L. mon sp. nov. and L. lahu sp. nov. mainly by the colour pattern of the larva (Fig.
Malaysia (Selangor), Indonesia (Sumatra), Thailand (Fig.
Holotype. Thailand • larva; Phetchaburi Prov., Kaeng Krachan Distr., Huai Mae Kamoei; 12°58'42"N, 99°34'55"E; 119 m; 24.ii.2018; leg. C. Suttinun; on slide; GBIFCH00980857;
Thailand • 1 larva; Prachuab Khiri Khan Prov., Kui Buri Distr., Huai Sam Rong; 100 m; 12°03'50"N, 99°37'39"E; 19.iv.2019; leg. C. Suttinun; on slide; GBIFCH01556160;
Larva. Following combination of characters differentiate L. mon sp. nov. from other species of the group batakorum: head and thorax dorsally mostly dark brown with indistinct brighter pattern; abdomen dorsally brown to dark brown, paler in middle area, segment V not much brighter; hind protoptera present, well developed.
Larva (Figs
Labiobaetis mon sp. nov., larva morphology: a. Hind leg; b. Hind claw; c. Section of hind femur; d. Section of hind tibia; e. Antennal base; f. Posterior margins of abdominal terga; g. Abdominal segment IX, posterior margin of tergum and sternum; h. Paraprocts; i, j. Tergalius IV; k. Left half of metanotum with hind protopteron. Scale bars: 50 µm (a); 20 µm (i); 10 µm (b–h, j).
Colouration
(Fig.
Antenna
(Fig.
Labrum
(Fig.
Right mandible
(Fig.
Left mandible
(Fig.
Both mandibles with lateral margins almost straight.
Hypopharynx and superlinguae
(Fig.
Maxilla
(Fig.
Labium
(Fig.
Hind protoptera
(Fig.
Legs
(Fig.
Abdominal terga
(Fig.
Abdominal sterna
(Fig.
Tergalii
(Fig.
Paraproct
(Fig.
Unknown.
The species is dedicated to the indigenous Mon people in Thailand.
Thailand (Fig.
Holotype. Thailand • larva; Tak Prov., Mueang Distr., Klong Lan Sang; 16°46'53"N, 99°01'16"E; 253 m; 25.xii.2017; leg. C. Suttinun; on slide; GBIFCH00596154;
Larva. Following combination of characters differentiate L. lahu sp. nov. from other species of the group batakorum: thorax and abdomen dorsally dark grey with yellowish pattern as in Fig.
Larva (Figs
Labiobaetis lahu sp. nov., larva morphology: a. Hind leg; b. Fore claw; c. Section of hind femur; d. Section of fore femur with femoral patch; e. Right half of metanotum with hind protopteron; f. Posterior margins of abdominal terga; g. Abdominal terga VIII and IX, posterolateral margin; h. Paraproct; i. Antennal base; j, k. Tergalius V. Scale bars: 50 µm (a, e); 20 µm (j); 10 µm (b–d, f–i).
Colouration
(Fig.
Antenna
(Fig.
Labrum
(Fig.
Right mandible
(Fig.
Left mandible
(Fig.
Both mandibles with lateral margins almost straight.
Hypopharynx
and superlinguae (Fig.
Maxilla
(Fig.
Labium
(Fig.
Hind protoptera
(Fig.
Legs
(Fig.
Abdominal terga
(Fig.
Abdominal sterna. Posterior margin of sterna: I–VI smooth, without spines; VII–IX with small, triangular spines.
Tergalii
(Fig.
Paraproct
(Fig.
Unknown.
The species is dedicated to the indigenous Lahu people in Thailand.
Thailand (Fig.
Diagnosis. Dorsal surface of labrum with submarginal row of simple setae, 1st and 2nd setae after submedian seta closely together; labial palp segment II with thumb-like protuberance; right mandible with pronounced hump between prostheca and mola, with thin setae basally at mola; distal margin of left mandible usually convex; maxillary palp segment II longer than segment I, usually bent; antennal scape without distolateral process; superlinguae strongly sclerotised along basal margin; glossae with robust setae along inner margin; vestigial hind protoptera; tergalii on abdominal segments II–VII; outer margin of femur with partial 2nd row of setae in proximal 1/2; tibia ventral margin with much longer seta distally at suture.
The following species are known from continental Southeast Asia: Labiobaetis numeratus (Müller-Liebenau, 1984), Labiobaetis tenasserimensis sp. nov., Labiobaetis angularis sp. nov., Labiobaetis tonsator sp. nov.
Holotype. Thailand • larva; Ratchaburi Prov., Suan Phueng Distr., Bo Klueng; 13°31'27"N, 99°14'39"E; 180 m; 25.xi.2018; leg. C. Suttinun; on slide; GBIFCH00763877;
Larva. Following combination of characters differentiate L. tenasserimensis sp. nov. from other species of the group numeratus: abdomen dorsally rather uniform brown; hypopharynx with well-developed medial tuft; labial palp segment II lobed, segment III rather short; spines at posterior margin of abdominal tergites wide, apically rounded, mostly not fused.
Larva (Figs
Labiobaetis tenasserimensis sp. nov., larva morphology: a. Middle leg; b. Outer margin of middle femur; c. Inner margin of middle femur; d. Outer margin of middle tibia; e. Inner margin of middle tibia; f. Middle claw; g. Posterior margins of abdominal terga; h. Paraproct; i, j. Tergalius IV; k. Antennal base. Scale bars: 50 µm (a); 20 µm (i); 10 µm (b–h, j, k).
Colouration
(Fig.
Antenna
(Fig.
Labrum
(Fig.
Right mandible
(Fig.
Left mandible
(Fig.
Both mandibles with lateral margins almost straight.
Hypopharynx and superlinguae
(Fig.
Maxilla
(Fig.
Labium
(Fig.
Hind protoptera vestigial.
Legs
(Fig.
Abdominal terga
(Fig.
Abdominal sterna
(Fig.
Tergalii
(Fig.
Paraproct
(Fig.
Unknown.
The most similar species is L. paranumeratus Kaltenbach & Gattolliat, 2019 from Sumatra. The following characters distinguish the larvae of both species (see
The species name is derived from the name of the Tenasserim mountain range where it was collected.
Thailand (Fig.
Holotype. Thailand • larva; Petchburi Prov., Kaeng Krachan Distr., Huai Mae Kamoei; 12°58'42"N, 99°34'55"E; 119 m; 24.ii.2018; leg. C. Suttinun; on slide; GBIFCH01223079;
Larva. Following combination of characters differentiate L. angularis sp. nov. from other species of the group numeratus: abdomen dorsally dark grey-brown, laterally off-white; left mandible with angular hump at margin between prostheca and mola; claw with 14–17 denticles; spines at posterior margin of abdominal tergites wide, apically rounded.
Larva (Figs
Labiobaetis angularis sp. nov., larva morphology: a. Labrum; b. Section of labrum, dorsal focus; c. Labium; d. Labial palp; e. Hypopharynx and superlinguae; f. Maxilla; g. Maxillary palp; h. Left mandible; i. Right mandible; j. Section of left mandible. Scale bars: 20 µm (c, e, f, h, i); 10 µm (a, b, d, g).
Labiobaetis angularis sp. nov., larva morphology: a. Middle leg; b. Middle tarsus and claw; c. Antennal base; d. Abdominal sterna VII–IX; e. Posterior margins of abdominal terga I, V, VIII; f. Paraproct; g. Left half of metanotum with vestigial hind protopteron; h–j. Tergalius IV. Scale bars: 50 µm (a); 20 µm (g, h); 10 µm (b–f, h–j).
Colouration
(Fig.
Antenna
(Fig.
Labrum
(Fig.
Right mandible
(Fig.
Left mandible
(Fig.
Both mandibles with lateral margins slightly convex.
Hypopharynx and superlinguae
(Fig.
Maxilla
(Fig.
Labium
(Fig.
Hind protoptera
(Fig.
Legs
(Fig.
Abdominal terga
(Fig.
Abdominal sterna
(Fig.
Tergalii
(Fig.
Paraproct
(Fig.
Unknown.
The Latin word angularis, meaning angular, refers to the unique angular hump at the margin between prostheca and mola of the left mandible.
Thailand (Fig.
Holotype. Thailand • larva; Patthalung Prov., Ton Sa Tor; 07°11'43"N, 100°04'18"E; 51 m; 12.iii.2016; leg. C. Suttinun; on slide; GBIFCH00829272;
Larva. Following combination of characters differentiate L. tonsator sp. nov. from other species of the group numeratus: abdomen dorsally grey-brown, segment II distomedially with dark grey-brown spot, segment V dark brown; pedicellus basally with dark brown hypodermal colouration; claw with 12–16 denticles; spines at posterior margin of abdominal tergites wide, apically rounded, mostly not fused.
Larva (Figs
Labiobaetis tonsator sp. nov., larva morphology: a. Middle leg; b. Outer margin of middle femur; c. Inner margin of middle femur; d. Inner margin of middle tibia; e. Outer margin of middle tibia; f. Fore tarsus and claw; g. Posterior margins of abdominal terga; h, i. Tergalius III; j. Paraproct; k. Antennal base. Scale bars: 50 µm (a); 20 µm (h); 10 µm (b–g, i–k).
Colouration
(Fig.
Antenna
(Fig.
Labrum
(Fig.
Right mandible
(Fig.
Left mandible
(Fig.
Both mandibles with lateral margins almost straight.
Hypopharynx and superlinguae
(Fig.
Maxilla
(Fig.
Labium
(Fig.
Hind protoptera vestigial.
Legs
(Fig.
Abdominal terga
(Fig.
Abdominal sterna . Posterior margin of sterna: I–VI smooth, without spines; VII–IX with wide triangular spines.
Tergalii
(Fig.
Paraproct
(Fig.
Unknown.
The species name is derived from the name of the village Ton Sa Tor, where it was collected (type locality).
Thailand (Fig.
Diagnosis. Labrum dorsal surface with submarginal row of feathered setae; labial palp segment II with large, thumb-like protuberance; antennal scape with well-developed distolateral process; hind protoptera well developed; tergalii present on abdominal segments I–VII.
The following species are known from continental Southeast Asia: Labiobaetis operosus (Müller-Liebenau, 1984), Labiobaetis cf. paraoperosus Kaltenbach & Gattolliat, 2018, Labiobaetis brao Kaltenbach, Garces & Gattolliat, 2022, Labiobaetis nisaratae sp. nov.
Thailand • 3 larvae; Ranong Prov., Mueang Distr., Punyaban Waterfall; 10°03'54"N, 98°40'13"E; 52 m; 19.vi.2018; leg. C. Suttinun; on slides; GBIFCH01223074, GBIFCH00596156;
Larva. Following combination of characters differentiate L. paraoperosus from other species of the group operosus: maxillary palp much longer than galea-lacinia, segment II with slight distolateral excavation; labial palp segment II with broad thumb-like, apically rounded distolateral protuberance; labial palp segment III oblong.
The larva has no morphological difference to L. paraoperosus from Indonesia (Sumatra), but there is a genetic distance of 12% (COI, K2P).
Thailand (Fig.
Holotype. Thailand • larva; Yasothon Prov., Kut Chum Distr., Huai Naso; 15°57'08"N, 104°18'12"E; 125 m; 11.08.2014; leg. B. Boonsoong; on slide; GBIFCH01221811;
Larva. Following combination of characters differentiate L. nisaratae sp. nov. from other species of the group operosus: abdomen dorsally brown, segments I–IV with paler areas as in Fig.
Larva (Figs
Labiobaetis nisaratae sp. nov., larva morphology: a. Fore leg; b. Section of fore femur; c. Section of fore tibia; d. Fore claw; e. Middle leg, femoral patch; f. Hind leg, femoral patch; g. Antennal base; h. Tergalius VII; i, j. Tergalius V; k. Posterior margins of abdominal terga; l. Paraproct. Scale bars: 50 µm (a, g–i); 10 µm (b–f, j–l).
Colouration
(Fig.
Antenna
(Fig.
Labrum
(Fig.
Right mandible
(Fig.
Left mandible
(Fig.
Both mandibles with lateral margins almost straight.
Hypopharynx and superlinguae
(Fig.
Maxilla
(Fig.
Labium
(Fig.
Hind protoptera
(Fig.
Legs
(Fig.
Abdominal terga
(Fig.
Abdominal sterna . Posterior margin of sterna: I–VII smooth, without spines; VIII and IX with triangular spines.
Tergalii
(Fig.
Paraproct
(Fig.
Male imago (Fig.
Colouration. Head and thorax beige with brown markings, turbinate eyes beige, shaft pink. Legs bluish. Wings and venation hyaline. Abdominal segments I–VI dorsally and ventrally bluish, segments VII, IX, and X off-white to beige, segment VIII light brown; abdominal segments dorsally with narrow brown bands at posterior margins. Cerci bluish.
Forewing . Pterostigma with seven cross-veins, first two reaching subcostal vein. Double intercalary veins mostly shorter than distance between corresponding main veins at wing margin.
Hind wing . Much smaller than forewing, with two longitudinal veins.
Genitalia. Basal segment of gonostylus (unistyliger) distally expanded at inner margin; segments I and II almost completely fused; segment I basally with protuberance at inner margin; small constriction at base of segment II; segment III slightly ovoid, with strong constriction at base, cross-section dimension slightly smaller than distal margin of segment II. Styliger plate between unistyligers well developed, distal margin straight.
The larva of L. nisaratae sp. nov. is very similar to L. operosus. The only clear difference is in the shape of labial palp segment II, where the distomedial protuberance in L. nisaratae sp. nov. is elongated, longer than the base of segment III (~1.1×). In L. operosus, the distomedial protuberance of segment II is shorter than the base of segment III (~0.8×) and stouter than in L. nisaratae sp. nov.
The name of this species is dedicated to Assoc. Prof. Dr. Nisarat Tungpairojwong (Department of Biology, Khon Kean University) for her outstanding contributions to the systematics of aquatic insects in Thailand.
Thailand (Fig.
Diagnosis. Labrum dorsal surface with submarginal row of clavate setae, submedian setae absent; left mandible without setae at mola apex; hind protoptera absent; tergalii usually on abdominal segments II–VII; distolateral process at scape absent or poorly developed;
The following species are known from continental Southeast Asia: Labiobaetis diffundus (Müller-Liebenau, 1984), Labiobaetis kui Kaltenbach, Garces & Gattolliat, 2022, Labiobaetis karen sp. nov., Labiobaetis septem sp. nov., Labiobaetis ranongensis sp. nov.
Holotype. Thailand • larva; Kanchanaburi Prov., Thong Pha Phum Distr., Pra Chum Mai; 14°34'58"N, 98°34'52"E; 269 m; 25.v.2017; leg. C. Suttinun; on slide; GBIFCH00763836;
Larva. Following combination of characters differentiate L. karen sp. nov. from other species of the group sumigarensis: thorax dorsally ochre, with dark brown distolateral dots on mesonotum; abdomen dorsally dark reddish-brown, paler in middle area, segment I ochre; femur with dark brown, triangular marking at inner, distomedial margin; tibia dark brown in subdistal area; scapus and pedicellus laterally with dark brown hypodermal colouration; labial palp segment III subrectangular with inner distal margin concave; paraproct distally not expanded.
Larva (Figs
Labiobaetis karen sp. nov., larva morphology: a. Fore leg; b. Outer margin of fore femur; c. Inner margin of fore femur; d. Section of fore tibia; e. Fore claw; f, g. Tergalius III; h. Posterior margins of abdominal terga; i. Paraproct; j. Antennal base. Scale bars: 50 µm (a); 20 µm (f); 10 µm (b–e, g–j).
Colouration
(Fig.
Antenna
(Fig.
Labrum
(Fig.
Right mandible
(Fig.
Left mandible
(Fig.
Both mandibles with lateral margins almost straight.
Hypopharynx and superlinguae
(Fig.
Maxilla
(Fig.
Labium
(Fig.
Hind protoptera absent.
Legs
(Fig.
Abdominal terga
(Fig.
Abdominal sterna . Posterior margin of sterna: I–VI smooth, without spines; VII–IX with triangular spines.
Tergalii
(Fig.
Paraproct
(Fig.
Unknown.
The species is dedicated to the indigenous Karen people in Thailand.
Thailand (Fig.
Holotype. Thailand • larva; Tak Prov., Mueang Distr., Klong Lan Sang; 16°46'53"N, 99°01'16"E; 253 m; 25.xii.2017; leg. C. Suttinun; on slide; GBIFCH01223075;
Thailand • 11 larvae; Tak Prov., Mueang Distr., Klong Lan Sang; 16°46'53"N, 99°01'16"E; 253 m; 25.xii.2017; leg. C. Suttinun; 3 on slides; GBIFCH00607174;
Larva. Following combination of characters differentiate L. septem sp. nov. from other species of the group sumigarensis: tergalii present on abdominal segments I–VII; abdomen dorsally uniform brown, or brown with crown-like pattern; femur with 9–15 spine-like setae at outer margin; tibia with row of short, apically rounded setae at outer margin, or bare with one medium, apically rounded seta distally; claw with 10–14 denticles.
Larva (Figs
Labiobaetis septem sp. nov., larva morphology: a. Fore leg (morphotype A); b. Fore leg (morphotype B); c. Section of fore femur (morphotype A); d. Section of fore tibia (morphotype A); e. Outer margin of tibia (morphotype B); f. Inner margin of fore tibia (morphotype B); g. Fore tarsus and claw. Scale bars: 50 µm (a, b); 10 µm (c–g).
Colouration
(Figs
Antenna
(Fig.
Labrum
(Fig.
Right mandible
(Fig.
Left mandible
(Fig.
Both mandibles with lateral margins almost straight.
Hypopharynx and superlinguae
(Fig.
Maxilla
(Fig.
Labium
(Fig.
Hind protoptera absent.
Legs
(Fig.
Abdominal terga
(Fig.
Abdominal sterna . Posterior margin of sterna: I–VI smooth, without spines; VII–IX with triangular spines.
Tergalii
(Fig.
Paraproct
(Fig.
The species appears with two different morphotypes, which partly share the same locations: based on our material, morphotype A is characterised by a dorsally rather uniform brown abdomen, a row of setae at outer margin of tibia, 10–15 setae at outer margin of femur, and a claw with 10 or 11 denticles; morphotype B has a crown-like pattern dorsally on the abdomen, outer margin of tibia almost bare, femur with nine or ten setae at outer margin, and claw with 12–14 denticles (see in discussion section). Holotype and paratypes correspond with morphotype A, specimens with morphotype B are mentioned as other material.
Unknown.
The Latin word septem, meaning seven, refers to the seven pairs of tergalii of this species. All other known species of the group sumigarensis have only six pairs of tergalii.
Thailand (Fig.
Holotype. Thailand • larva; Ranong Prov., Mueang Distr., Huai Nam Sai; 09°43'25"N, 98°36'29"E; 50 m; 20.vi.2018; leg. C. Suttinun; on slide; GBIFCH00980864;
Larva. Following combination of characters differentiate L. ranongensis sp. nov. from other species of the group sumigarensis: abdomen dorsally yellow-brown, basal parts of terga paler, posterior parts with a darker, slightly crown-like marking; labial palp segment III subrectangular, segment II with thumb-like protuberance with straight distal margin and rounded lateral margin; maxillary palp longer than galea-lacinia, terminal segment with well-developed, distolateral excavation; left mandible with margin between prostheca and mola straight, with minute marginal and submarginal denticles; abdominal tergites II–IX with triangular, sharply pointed spines on posterior margins; paraproct distally not expanded, with ~28 spines.
Larva (Figs
Labiobaetis ranongensis sp. nov., larva morphology: a. Hind leg; b. Section of hind femur; c. Section of hind tibia; d. Hind tarsus and claw; e. Antennal base; f. Posterior margins of abdominal terga; g. Posterior margins of abdominal segment IX; h. Paraprocts; i, j. Tergalius V. Scale bars: 50 µm (a); 20 µm (g); 10 µm (b–f, h–j).
Colouration
(Fig.
Antenna
(Fig.
Labrum
(Fig.
Right mandible
(Fig.
Left mandible
(Fig.
Both mandibles with lateral margins almost straight.
Hypopharynx and superlinguae
(Fig.
Maxilla
(Fig.
Labium
(Fig.
Hind protoptera absent.
Legs
(Fig.
Abdominal terga
(Fig.
Abdominal sterna
(Fig.
Tergalii
(Fig.
Paraproct
(Fig.
Unknown.
The species name refers to Ranong Prov., where the type locality is located.
Thailand (Fig.
| 1 | Labrum dorsally with arc of simple setae | 2 |
| – | Labrum dorsally with arc of feathered or clavate setae | 8 |
| 2(1) | With well-developed hind protoptera (batakorum species group) | 3 |
| – | With minute hind protoptera (numeratus species group) | 5 |
| 3(2) | Abdomen dorsally rather uniform brown to dark brown, paler in middle area, tergum V not much brighter (Fig. |
L. mon sp. nov. |
| – | Abdomen dorsally with lively pattern, tergum V much brighter than neighbouring terga (Figs |
4 |
| 4(3) | Abdomen dorsally dark grey with yellowish pattern as in Fig. |
L. lahu sp. nov. |
| – | Abdomen dorsally grey-brown, laterally whitish with black markings, terga V and X brighter (Fig. |
L. multus |
| 5(2) | Left mandible with angular hump at margin between prostheca and mola (Fig. |
L. angularis sp. nov. |
| – | Left mandible without angular hump at margin between prostheca and mola | 6 |
| 6(5) | Abdomen dorsally grey-brown, tergum II laterally darker with dark brown, roundish, distomedial marking, tergum V dark brown | L. tonsator sp. nov. |
| – | Abdomen dorsally rather uniform brown | 7 |
| 7(6) | Labial palp segment III approx. as long as wide at base ( |
L. numeratus |
| – | Labial palp segment III rather short (length 0.8× width at base; Fig. |
L. tenasserimensis sp. nov. |
| 8(1) | Labrum dorsally with arc of feathered setae (operosus species group) | 9 |
| – | Labrum dorsally with arc of clavate setae (sumigarensis species group) | 12 |
| 9(8) | Labial palp segment II with broad thumb-like (lobed), distolateral protuberance, segment III oblong; maxillary palp segment II with slightly developed distolateral excavation ( |
L. cf. paraoperosus |
| – | Labial palp segment II with rather narrow or elongate thumb-like, distomedial protuberance (Figs |
10 |
| 10(9) | Labial palp segment II with thumb-like, distomedial protuberance, slightly bent distad; paraproct at inner, proximal margin with additional submarginal row of minute spines ( |
L. brao |
| – | Labial palp segment II with thumb-like, distomedial protuberance, directed laterally, proximal margin of protuberance slightly concave (Figs |
11 |
| 11(10) | Distomedial protuberance of labial palp segment II shorter than base of segment III (~0.8×) (Fig. |
L. operosus |
| – | Distomedial protuberance of labial palp segment II elongated, longer than base of segment III (~1.2×) (Fig. |
L. nisaratae sp. nov. |
| 12(8) | Tergalii on abdominal segments I–VII | L. septem sp. nov. |
| – | Tergalii on abdominal segments II–VII | 13 |
| 13(12) | Antennal scape with reduced, distolateral process ( |
L. diffundus |
| – | Antennal scape without distolateral process | 14 |
| 14(13) | Thorax dorsally ochre with pronounced, dark brown, distolateral markings on mesonotum; abdomen dorsally dark reddish-brown, slightly paler in middle area, tergum I ochre; abdomen ventrally mainly beige, laterally dark reddish-brown (Fig. |
L. karen sp. nov. |
| – | Thorax dorsally pale brown, no pronounced distolateral markings on mesonotum; abdomen dorsally pale brown, ventrally rather uniform beige | 15 |
| 15(14) | Posterior margin of abdominal terga with triangular spines, sometimes spaced; abdominal terga dorsally rather uniform brown ( |
L. kui |
| – | Posterior margin of abdominal terga with triangular, sharply pointed spines (Fig. |
L. ranongensis sp. nov. |
Habitats: a. Labiobaetis multus; b. Labiobaetis mon sp. nov. (TL) and angularis sp. nov. (TL); c. Labiobaetis lahu sp. nov.; d. Labiobaetis tenasserimensis sp. nov. (TL); e. Labiobaetis tonsator sp. nov. (TL); f. Labiobaetis nisaratae sp. nov. and L. karen sp. nov. (TL); g. Labiobaetis septem sp. nov.; h. Labiobaetis ranongensis sp. nov. (TL). TL: type locality.
For the assignment of the new species to Labiobaetis we refer to
The morphological species groups within Labiobaetis are primarily a working tool, but could also serve as a basis for future studies on the generic or subgeneric delimitation and phylogeny of this genus. Further investigation including the analysis of nuclear gene sequences may prove that some are natural groups.
We obtained COI sequences from all Labiobaetis species reported from Thailand, including the nine newly described species (Table
The genetic distances between Labiobaetis species in Thailand (10–25%, K2P) is rather high, which is in line with the interspecific genetic distances found in Indonesia (11–24%;
Interestingly, the new species Labiobaetis septem sp. nov. presents two different morphotypes. Morphotype A has a dorsally rather uniform brown abdomen (Fig.
Specimens were collected primarily from headwater and urban streams across a range of altitudes (10–810 m a.s.l.). Most streams were situated in forested areas with partially closed canopies, while a few were in urban environments with riparian vegetation and partially covered canopies. In headwater streams, the substrate was predominantly composed of gravel and sand. In contrast, urban streams had sandy substrates, with Hydrilla verticillata (L.f.) Royle (commonly known as Water Thyme) as dominant aquatic vegetation. The larvae were found in the flowing water.
The genus Labiobaetis was the most dominant and widely distributed baetid genus across all sampling sites in Thailand. It was present in every locality surveyed and consistently comprised multiple species at each site. In some cases, up to 100 specimens were collected per sampling, indicating that Labiobaetis is the most abundant genus within the family Baetidae in Thailand.
Considering the general mega-biodiversity in Thailand, and the obvious richness of Labiobaetis in this region, we have to expect many more new species with further studies in the future, despite the high collection efforts already done in the past years.
We sincerely thank Chiang Mai University, National Research Council of Thailand (NRCT) and the Centre of Excellence on Biodiversity (BDC) Office of Higher Education Commission (BDC-PG2-161004) for the support of field work. Further, we are grateful to Céline Stoffel and Marion Podolak (Naturéum, Muséum Cantonale des Sciences Naturelles,
The authors have declared that no competing interests exist.
We followed all guidelines of the Animal Ethics Committee of Kasetsart University (approval no. ACKU61-SCI-028, ACKU63-SCI-006, ACKU66-SCI-017) for collecting the mayfly specimens.
No use of AI was reported.
National Research Council of Thailand (NRCT) contract Number N42A660904.
Formal analysis: CS. Funding acquisition: BB, CS. Investigation: TK, BB, CS. Project administration: BB. Validation: CS. Visualization: TK. Writing – original draft: TK. Writing – review and editing: CS, JLG, BB.
Thomas Kaltenbach https://orcid.org/0000-0001-8052-0388
Jean-Luc Gattolliat https://orcid.org/0000-0001-5873-5083
Boonsatien Boonsoong https://orcid.org/0000-0002-8166-0021
Chanaporn Suttinun https://orcid.org/0000-0002-0380-1554
All of the data that support the findings of this study are available in the main text or Supplementary Information.
Labiobaetis species of Thailand, genetic distances (COI, K2P)
Data type: xls