Research Article |
Corresponding author: Boonsatien Boonsoong ( fscibtb@ku.ac.th ) Academic editor: Lyndall Pereira-da-Conceicoa
© 2021 Sedtawut Kwanboon, Michel Sartori, Boonsatien Boonsoong.
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:
Kwanboon S, Sartori M, Boonsoong B (2021) Behningiidae and Potamanthidae (Insecta, Ephemeroptera) in Thailand. ZooKeys 1067: 57-82. https://doi.org/10.3897/zookeys.1067.72779
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The aim of this study is to review the family Behningiidae and Potamanthidae in Thailand. Two genera and three species of Behningiidae are recognised: Protobehningia merga Peters & Gillies, 1991, Behningia baei McCafferty & Jacobus, 2006, and Behningia nujiangensis Zhou & Bisset, 2019, which is newly reported from Thailand. The egg structure of B. nujiangensis is described for the first time using scanning electron microscopy. The larva of P. merga is redescribed and its distribution is expended northward with a new record from Chiang Mai province. Two genera and five species of Potamanthidae are identified: Potamanthus formosus Eaton, 1892, Rhoenanthus magnificus Ulmer, 1920 (new record for Thailand), Rhoenanthus obscurus Navás, 1922, Rhoenanthus distafurcus Bae & McCafferty, 1991, and Rhoenanthus speciosus Eaton, 1881. Our morphological evidence is supported by COI data for the family Potamanthidae. Diagnostic characters, distributions, and keys are presented for the larvae of all known species of Thai behningiid and potamanthid mayflies.
Biodiversity, COI, egg, mayfly, new record
The Behningiidae is a small mayfly family represented by three extant genera (Behningia Lestage, 1930, Dolania Edmunds & Traver, 1959, and Protobehningia Tshernova, 1960) and one fossil genus (Archaeobehningia Tshernova, 1977) (
The family Potamanthidae is widely distributed throughout the Holarctic and Oriental regions and accounts for 25 species worldwide. In Southeast Asia, seven species in two genera and four subgenera have been reported (
In this study, we review the species of Behningiidae and Potamanthidae in Thailand, and we provide the first records of B. nujiangensis Zhou & Bisset, 2019 and R. magnificus Ulmer, 1920. We also redescribe the larva of P. merga, and we present the first description of the egg structure of B. nujiangensis. A distribution map of Thai behningiid and potamanthid mayflies is also provided.
The specimens were collected from streams and rivers in Thailand and were preserved in absolute ethanol. Measurements (in mm) and photographs were taken using a Nikon SMZ800 and ZEISS Stemi 305 stereoscopic microscope. For scanning electron microscopy (SEM), specimens (head, legs, labrum, labium, labial palp, glossa, paraglossa, and eggs) were dried in a critical point dryer (CPD7501) and coated with gold (Sputter Coater SC7620). The specimens were observed and photographed with an FEI Quanta 450 SEM. The final plates were prepared with Adobe Photoshop CC 2020. The material is deposited in the collection of the Zoological Museum at Kasetsart University in Bangkok, Thailand (ZMKU). The distribution map was constructed using the Simple Mapper website (http://www.simplemappr.net) and GPS coordinates.
Each specimen was dissected for DNA extraction using a genomic DNA purification kit (NucleoSpin, Macherey-Nagel, Germany) following the manufacturer’s protocol. A fragment of the mitochondrial cytochrome oxidase I (COI) was amplified using the primers LCO1490 and HCO2198 (
Species | Code | Collection locality | Collector | Date | GenBank accession number |
---|---|---|---|---|---|
Protobehningia merga | PM01CM | Chiang Mai | B. Boonsoong | 13 Nov 2020 | MW792224 |
Potamanthus formosus | PF01NA | Nan | B. Boonsoong | 28 Nov 2020 | MZ453438 |
PF02KN | Kanchanaburi | S. Kwanboon | 11 Jul 2019 | MZ453439 | |
PF03CR | Chiang Rai | S. Kwanboon | 6 Mar 2021 | MZ436659 | |
PF04CR | Chiang Rai | S. Kwanboon | 5 Mar 2021 | MZ436660 | |
Rhoenanthus magnificus | RM01NA | Nan | S. Kwanboon | 10 Mar 2018 | MZ436661 |
RM04NA | Nan | B. Boonsoong | 28 Nov 2020 | MZ436662 | |
RM05CR | Chiang Rai | S. Kwanboon | 6 Mar 2021 | MZ436663 | |
RM06CR | Chiang Rai | S. Kwanboon | 7 Mar 2021 | MZ436664 | |
R. obscurus | RO02FCM | Chiang Mai | S. Kwanboon | 15 Nov 2020 | MZ436665 |
ROO7CM | Chiang Mai | S. Kwanboon | 15 Nov 2020 | MZ436666 | |
R. distafurcus | RD01NA | Nan | B. Boonsoong | 28 Nov 2020 | MZ436667 |
RD02NA | Nan | B. Boonsoong | 28 Nov 2020 | MZ436668 | |
RD03KN | Kanchanaburi | B. Boonsoong | 15 Oct 2015 | MZ436669 | |
RD04RB | Ratchaburi | B. Boonsoong | 19 Apr 2016 | MZ436670 |
Order Ephemeroptera Hyatt & Arms, 1891
Family Behningiidae Motas & Bacesco, 1937
None.
The larvae of Behningia baei McCafferty & Jacobus, 2006 can be distinguished from other Behningia species based on the following characteristics: i) labrum deeply emarginate in a V or U shape at anteromedian margin, ii) labial palp I without concavity on outer margin, iii) labial palp II less than 50% length of labial palp III, iv) tarsus of foreleg as long as tibia and v) coxa of hind leg less than 60% as long as femur.
Phitsanulok province.
The larvae of B. baei were originally described by
2 mature larvae, Thailand, Chiang Mai province, Mae Tang district, Tard Luang Waterfall, 19°01'27.5"N, 98°51'17.1"E, 18.IX.2011, P. Sritipsak leg. deposited in ZMKU.
The larvae of Behningia nujiangensis Zhou & Bisset, 2019 can be separated from those of other Behningia species based on the following characteristics: i) labrum shallowly emarginate at anteromedian margin (Fig.
Egg (dissected from mature larva). Length 1.62–1.73 mm, width 1.09–1.26 mm (n = 13); oval (Fig.
Chiang Mai province.
The larvae of B. nujiangensis were originally described by
2 larvae, deposited in ZMKU, Thailand, Chiang Mai province, Mae Chaem district, Mae Chaem river, 18°30'46.0"N 98°21'22.6"E, 475 m, 5.X.2019, B. Boonsoong leg., 1 larva, same data, 13.XI.2020, B. Boonsoong leg. (ZMKU).
Larva (in alcohol, Fig.
Head. Anterior margin not projecting, front with densely short goldish setae standing out on the head (Fig.
Thorax. Colour pale yellowish. Forelegs flattened, with large broad coxae, flat femur, small claws (Fig.
Abdomen. Similar in colour to head and thorax, abdominal segments elongated and convex, with short straight setae at lateral margin, lateral margin of abdominal segment I–IX with flat projections spine-like in shape. Gill present on segment I–VII, plumose shape, first gill filament single (Fig.
The larvae of Protobehningia merga Peters & Gillies 1991 can be distinguished from those of P. asiatica based on the following characteristics: i) glossae and paraglossae with more than 20 setae on the ventral surface, ii) maxillary palp segment II 2/3 the length of segment I, each maxillary palp segment completely divided, but segments II and III indistinct (
Kanchanaburi and Chiang Mai provinces.
In general, the larvae of behningiid mayflies are rarely collected. In this study, the larvae (middle instar) were found in October (turbidity from flooding, Fig.
Only two species of Protobehningia are known in the world: Protobehningia asiatica Tshernova & Bajkova 1960 and Protobehningia merga Peters & Gillies 1991.
(adapted from
1 | Tarsi of forelegs not fused to tibiae (Fig. |
Behningia , 2 |
– | Tarsi of forelegs fused to tibiae (Fig. |
Protobehningia , P. merga |
2 | Medio-anterior emargination of labrum deep ( |
Behningia baei |
– | Medio-anterior emargination of labrum very shallow (Fig. |
Behningia nujiangensis |
Genus Potamanthus Pictet, 1843
SubgenusPotamanthodes Ulmer, 1920
1 larva, Thailand, Chanthaburi province, Makham district, Ban Pa Rim Tarn homestay, 12°51'00.0"N, 102°12'17.1"E, 5.X.2019, B. Boonsoong leg. (ZMKU); 2 larvae, Kanchanaburi province, Huai Pak Kok, 14°39'34.4"N, 98°32'02.3"E, 175 m, 11.VII.2019, S. Kwanboon leg. (ZMKU); 2 larvae, Chiang Rai province, Huai Kang Pla waterfall, 20°05'21.6"N, 99°46'47.8"E, 519 m, 5.III.2021, S. Kwanboon leg. (ZMKU); 4 larvae, Chiang Rai province, Klong Mae Salong, 20°09'52.0"N, 99°40'06.8"E, 6.III.2021, S. Kwanboon leg. (ZMKU); 1 larva, Nan province, Ban Ratsadonsamakkhi, 18°52'23.4"N, 100°49'54.1"E, 59 m, 28.XI.2020, B. Boonsoong leg.
The larvae of Potamanthus formosus Eaton, 1892 can be distinguished from those of other Potamanthus (Potamanthodes) species based on the following characteristics: i) dorsal forefemora with simple stout setae (Fig.
Chanthaburi, Kanchanaburi, Nan, and Chiang Rai provinces.
The adult of P. formosus was described by
SubgenusPotamanthindus Lestage, 1930
5 larvae, Thailand, Chiang Mai province, Chiang Dao, Mae Na, 19°19'13.08"N, 98°53'25.98"E, 742 m, 11.III.2016, B. Boonsoong leg. (ZMKU); 8 larvae, Loel province, Nam Thob ranger station, 17°15'36.5"N, 101°34'52.9"E, 338 m, 20.III.2016, B. Boonsoong leg. (ZMKU); 1 larva, Nan province, Bo Kluea district, Sapan waterfall, 19°11'25.8"N, 101°11'56.3"E, 800 m, 28.XI.2020, B. Boonsoong leg; 2 larvae, Nan province, Bo Kluea district, Lamer resort, 19°09'08.8"N, 101°09'17.0"E, 28.XI.2020, S. Kwanboon leg; 3 larvae, Nan province, Bo Kluea district, Mae Nam Wa stream, 19°16'22.6" N 101°10'48.2" E, 848 m, 26.XI.2019, B. Boonsoong leg; 7 larvae, Chiang Rai province, Mueang district, Mae Kon stream, 19°51'46.1"N, 99°39'04.7"E, 534 m, 6.III.2021, S. Kwanboon leg; 2 larvae, Chiang Rai province, Mueang district, Mae Kon stream, Pong Phrabat waterfall, 20°00'41.8"N, 99°48'15.1"E, 470 m, 7.III.2021, S. Kwanboon leg.
The larvae of Rhoenanthus magnificus (Fig.
Chiang Mai, Chiang Rai, Loei, and Nan provinces.
The larva of R. magnificus was originally described by
1 female imago (reared) and 1 male imago (reared), Thailand, Chiang Mai province, Mae Ping river, Elely Cafe, 19°04'08.4"N, 98°56'28.8"E, 15.XI.2020, S. Kwanboon leg. (ZMKU).
The larvae of Rhoenanthus obscurus can be distinguished from those of other Rhoenanthus (Potamanthindus) species based on the following characteristics: i) medium-sized body (12–17 mm), ii) mandibular tusks arched inward about 28° (angle measurement as shown in Fig.
Chiang Mai province.
The larva of R. obscurus was originally described by
1 larva, Thailand, Kanchanaburi province, Sai Yok district, Pueng Wahn Resort, 14°12'08.9"N, 99°03'36.0"E, 15.X.2015, B. Boonsoong leg; 2 larvae, Ratchaburi province, Suan Phueng district, Pha Chi river, 13°30'57.3"N, 99°20'40.1"E, 19.IX.2016, B. Boonsoong leg; 1 larva, Nan province, Bo Kluea district, Sapan waterfall, 19°11'25.8"N, 101°11'56.3”E, 800 m, 28.XI.2020, B. Boonsoong leg.
The larvae of Rhoennanthus distafurcus can be distinguished from those of other Rhoenanthus (Rhoenanthus) species based on the following characteristics: i) subapical spine of the mandibular tusk well developed laterally (Fig.
Kanchanaburi, Ratchaburi, and Nan provinces.
5 larvae, Thailand, Narathiwat province, Klong Aika Ding stream, 5°47'45.9"N, 101°50'05.5"E, 22.IV.2018, B. Boonsoong leg.
The larvae of Rhoenanthus speciosus can be distinguished from those of other Rhoenanthus (Rhoenanthus) species based on the following characteristics: i) lateral subapical spine of the mandibular tusk present (Fig.
Egg (dissected from mature larva). Oval; with two large conical polar caps, (Fig.
Narathiwat and Songkla provinces.
The larvae of Rhoenanthus speciosus were reported by
(adapted from
1 | Mandibular tusks subequal to, or longer than 1/2 length of head (Fig. |
Rhoenanthus, 2 |
– | Mandibular tusks shorter than 1/2 length of head (Fig. |
Potamanthus,subgenus Potamanthodes, P. formosus |
2 | Mandibular tusks with lateral subapical spine, appearing apically forked | subgenusRhoenanthus, 3 |
– | Mandibular tusks without lateral subapical spine, not appearing apically forked | subgenusPotamanthindus, 4 |
3 | Mandibular tusks with large lateral subapical spine with 16–20 medial rounded setae (Fig. |
R. distafurcus |
– | Mandibular tusks with small lateral subapical spine, without medial rounded setae (Fig. |
R. speciosus |
4 | Mandibular tusks strongly convergent and abruptly curved inward about 33–34° (Fig. |
R. magnificus |
– | Mandibular tusks strongly convergent and abruptly curved inward about 28° (Fig. |
R. obscurus |
The partial sequence of the mitochondrial COI gene (658 bp) of P. merga (MW792224) found in Thailand was analysed and compared with the sequence of Dolania americana (BIT011-04) from BOLD. However, there is no available sequence for the genus Behningia. The intergeneric genetic distance between these two genera was 22.39%, as determined by the Kimura 2-parameter (K2P) model. For Potamanthidae, the phylogenetic tree of the ML analysis is shown in Figure
Pairwise genetic distances (COI) between species of Potamanthidae using the Kimura 2-parameter.
1 | 2 | 3 | 4 | ||
---|---|---|---|---|---|
1 | Potamanthus formosus | – | |||
2 | Rhoenanthus obscurus | 0.171–0.190 | – | ||
3 | Rhoenanthus magnificus | 0.181–0.212 | 0.125–0.158 | – | |
4 | Rhoenanthus distafurcus | 0.177–0.198 | 0.171–0.196 | 0.182–0.210 | – |
The discovery of an additional species of Behningia in Thailand reveals the high diversity of the behningiid mayflies in the country. The presence of B. nujiangensis was confirmed based on morphological evidence according to
The presence of R. magnificus in Thailand was confirmed based on the morphological characters proposed by
The larvae of Behningiidae are restricted to fine sandy habitats (
This research is funded by Kasetsart University through the Graduate School Fellowship Program. We are most grateful to Assistant Professor Dr Akekawat Vitheepradit and Mr Prabsuek Sritipsak (Department of Entomology, Faculty of Agriculture, Kasetsart University) for kindly provide specimens for this study. This research has been supported by the Centre of Excellence on Biodiversity (BDC) Office of Higher Education Commission (BDC-PG2-161004). This research was approved by the Institutional Animal Care and Use Committee, Faculty of Science, Kasetsart University, Thailand under Project number ACKU61-SCI-028. We would like to thank the Department of Zoology and the Faculty of Science at Kasetsart University in Bangkok for their assistance and use of their facilities.