Research Article |
Corresponding author: Boonsatien Boonsoong ( fscibtb@ku.ac.th ) Academic editor: Ben Price
© 2019 Boonsatien Boonsoong, Michel Sartori.
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:
Boonsoong B, Sartori M (2019) Review and integrative taxonomy of the genus Prosopistoma Latreille, 1833 (Ephemeroptera, Prosopistomatidae) in Thailand, with description of a new species. ZooKeys 825: 123-144. https://doi.org/10.3897/zookeys.825.32443
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Three species of the genus Prosopistoma Latreille, 1833 (Prosopistomatidae) are currently reported from Thailand. A new species, Prosopistoma carinatum sp. n., is described here based on specimens from western and southern Thailand. The new species can be easily distinguished from the other members of Prosopistoma by the following combination of characteristics: (i) the presence of two ridged longitudinal lines on each side of its carapace, (ii) antenna 7-segmented, (iii) a strongly convex carapace and (iv) nine pectinate setae on the ventral margin of the fore tibiae. A comparison between the key characteristics of P. carinatum sp. n. and the known Thai species is provided. Results of analysis of the mitochondrial cytochrome oxidase I (COI) gene (658 bp) of three species, as well as the distribution of the Thai species, are also discussed.
Prosopistoma carinatum , Prosopistoma annamense , Prosopistoma sinense , COI, mayfly
Prosopistomatidae (Ephemeroptera) is a monogeneric family, represented by the genus Prosopistoma, which was originally established by
In Thailand, only the larvae of Prosopistoma annamense Soldán & Braasch, 1984, P. sinense Tong & Dudgeon, 2000, and P. wouterae Lieftinck, 1932 are known (
The prosopistomatid mayfly larvae were collected from streams and rivers in northern, north-eastern, eastern, central, western, and southern Thailand from 2013 to 2018. Measurements (given in mm) and photographs were taken using a Visionary LK System (Dun, Incorporated, USA). All drawings were made with the aid of a camera lucida attached to a compound microscope. For Scanning electron microscopy (SEM), specimens (head, carapace, sternal plate, foreleg) were dried in a Critical Point Drier (CPD7501) and coated with gold (10 nm, Sputter Coater SC7620). SEM photographs were observed by a FEI Quanta 450 SEM. Final plates were prepared with Adobe Photoshop CC 2017. The material is deposited in the collection of the Zoological Museum at Kasetsart University in Bangkok, Thailand (ZMKU) and at the Museum of Zoology in Lausanne, Switzerland (MZL). The distribution map was generated via the Simple Mapper website (http://www.simplemappr.net) using GPS coordinates.
The collected specimens were fixed in absolute ethanol and preserved under refrigeration for description and DNA extraction. Details of the specimens from the three species used for the DNA experiment are shown in Table
Species | Code | Collection locality | Collector | Date | GenBank accession number |
---|---|---|---|---|---|
P. annamense | PA-01-KN | Kanchanaburi | B. Boonsoong | 15 Oct 2015 | MK285321 |
PA-12-RT | Ratchaburi | B. Boonsoong | 13 Feb 2016 | MK285322 | |
PA-13-LE | Loei | B. Boonsoong | 19 Mar 2016 | MK285323 | |
PA-17-NN | Nakhon Nayok | B. Boonsoong | 25 Feb 2017 | MK285324 | |
PA-18-CT | Chantaburi | B. Boonsoong | 5 Jul 2018 | MK285325 | |
P. carinatum sp. n. | PC-01-KN | Kanchanaburi | B. Boonsoong | 26 Apr 2014 | MK285326 |
PC-02-KN | Kanchanaburi | B. Boonsoong | 26 Apr 2014 | MK285327 | |
P. sinense | PS-01-KN | Kanchanaburi | B. Boonsoong | 20 Feb 2016 | MK285328 |
PS-02-KN | Kanchanaburi | B. Boonsoong | 20 Feb 2016 | MK285329 | |
PS-03-CM | Chaing Mai | B. Boonsoong | 11 Mar 2017 | MK285330 |
C central;
m meter;
N northern;
NE north-eastern;
W western.
Prosopistoma
annamense
Soldán & Braasch, 1984: 370–376, figs. 2, 4, 6, 8, 13, 14. (orig.); Barber-James, 2009: 153–154, Table
THAILAND; 1 larva, Chanthaburi province, Klong Kla Seu Yai, Khao Kitchakut, 12°52'35.94"N, 102°05'48.3"E, 40 m, 7.II.2013, 2 larvae, same place, 6.VII.2018, B Boonsoong leg. (ZMKU); 1 larva, Nakhon Nayok province, Wang Ta Krai, 14°19'35.9"N, 101°18'05.9"E, 65 m, 24.III.2013, B Boonsoong leg. (ZMKU); 5 larvae, Kanchanaburi province, Thong Pha Phum, Phu Iyara Resort, 14°37'34.4"N, 98°34'17.0"E, 207 m, 6.IV.2013, B Boonsoong leg. (ZMKU), 1 larva same data (MZL, GBIFCH00657966); 1 larva, Ban Pra Chum Mai, 14°35'01.4"N, 98°34'54.3"E, 233 m, 26.IV.2014, B Boonsoong leg. (ZMKU); 5 larvae, Kanchanaburi province, Pung Wan Resort, 14°12'20"N, 98°03'36"E, 28 m, 14.X.2015, B Boonsoong leg. (ZMKU), 1 larva same data (MZL, GBIFCH00657965); 3 larvae, Loei province, Ban Non Pattana, 17°06'24"N, 101°28'44"E, 530 m, 1.II.2016, B Boonsoong leg. (ZMKU); 3 larvae, Ratchaburi province, Kang Som Maew, Suan Phueng, 13°24'27.6"N, 99°16'51.3"E, 206 m, 13.II.2016, B Boonsoong leg. (ZMKU); 6 larvae, Loei province, Ban Non Pattana, 17°06'24"N, 101°28'44"E, 530 m, 19.III.2016, B Boonsoong leg. (ZMKU); 2 larvae, Nakhon Nayok province, Wang Ta Krai, 14°20'9.42"N, 101°18'22.5"E, 67 m, 25.II.2017, B Boonsoong leg. (ZMKU); 2 larvae, Chanthaburi province, Klong Sa Tor Bon, 12°43'12.42"N, 102°23'19.26"E, 115 m, 5.VII.2018, B Boonsoong leg. (ZMKU).
The larvae of P. annamense can be distinguished from those of other Oriental congeners by (i) apex of the inner margin of the fore tibiae, with 4–6 serrated pectinate spines (Fig.
Loei province (NE), Nakhon Nayok province (C), Chanthaburi province (E), Kanchanaburi province (W), Ratchaburi province (W).
The larvae of P. annamense were originally described by
Uncorrected pairwise genetic distances (COI) between selected prosopistomatid and baetiscid species, using the Kimura 2-parameter.
B. laurentina | P. annamense | P. carinatum sp. n. | P. sinense | |
---|---|---|---|---|
B. laurentina | – | – | – | – |
P. annamense | 0.32 | – | – | – |
P. carinatum sp. n. | 0.23 | 0.31 | – | – |
P. sinense | 0.34 | 0.33 | 0.35 | – |
Prosopistoma
sinense
Tong & Dudgeon, 2000: 122–128, figs. 1–14. (orig.); Barber-James, 2009: 153–154, Table
THAILAND; 2 larvae, Kanchanaburi province, Thong Pha Phum, Pussaduklang Ranger Station, 14°33'13"N, 98°34'17"E, 317 m, 20.II.2016, B Boonsoong leg. (ZMKU); 2 larvae, Kanchanaburi province, Thong Pha Phum, Pak Kok stream, 14°39'32.1"N, 98°31'59.2"E, 161 m, 20.II.2016, B Boonsoong leg. (ZMKU), 1 larva same data (MZL, GBIFCH00657969); 4 larvae, Chiang Mai province, Chiang Dao, Mae Na, 19°19'13.08"N, 98°53'25.98"E, 742 m, 11.III.2017, B Boonsoong leg. (ZMKU).
The larvae of Prosopisoma sinense can be distinguished from those of other congeners by (i) antennae with 4–5 segments, (ii) antenna segment III much longer than the combined length of segments IV–V (iii) and apex of the ventral margin of the fore-tibia with 6–8 serrated spines (Fig.
Kanchanaburi province (W), Chiang Mai province (N).
The larvae of P. sinense were originally described by
Prosopistoma
wouterae
Lieftinck, 1932: 44–55, pls 1–2. (orig.); Lafon, 1952: 433 (table); Peters, 1967: 211–213, figs 2, 8, 10, 16, 26 (redescription); Soldán & Braasch, 1984; 374–375 (key); Barber-James, 2009: 153–154, Table
None.
The larvae of P. wouterae can be distinguished from those of other congeners by the combination of the following characteristics: (i) apical segment of the maxillary palp shorter than 1/2 the length of segment II, (ii) mesonotum with five dark-brown patches connected by brown narrow stripes, (iii) apex of the ventral margin of the fore-tibiae with eight serrated spines and (iv) posterolateral spine of segments VII and VIII parallel to or bent inwards the body axis (
Songkhla province (S).
Prosopistoma wouterae was originally described by
Holotype. THAILAND; Holotype, 1 mature larva, Kanchanaburi province, Thong Pha Phum, Ban Pra Chum Mai, 14°35'01.4"N, 98°34'54.3"E, 233 m, 11.IV.2015, B Boonsoong leg. (ZMKU); Paratypes. 1 larva same data as holotype (MZL, GBIFCH00657926); 6 larvae same data as holotype (ZMKU); 1 mature larva, 8 larvae, Kanchanaburi province, Ban Pra Chum Mai, 14°35'01.4"N, 98°34'54.3"E, 233 m, 26.IV.2016, B Boonsoong leg. (ZMKU).
THAILAND; 1 larva, Nakhon Si Thammarat province, Lan Sa Ka, Khao Luang water fall, 8°28'08.30"N, 99°46'14.20"E, 533 m, 2.VII.2016, B Boonsoong leg. (ZMKU); 1 larva, Narathiwat province, Waeng district, Ai Sae, 5°47'45.9"N, 101°50'5.46"E, 64 m, 21.IV.2018, B Boonsoong leg. (ZMKU).
Kanchanaburi province (W), Nakhon Si Thammarat province (S), Narathiwat province (S).
The larvae of Prosopistoma carinatum sp. n. can be distinguished from those of other species by the combination of the following characteristics (Table
Larval characters of P. carinatum sp. n. compared with Thai known species (
P. annamense | P. sinense | P. wouterae | P. carinatum sp. n. | |
---|---|---|---|---|
Distribution | Vietnam, Thailand | China, Thailand | Java, Sumatra, Thailand | Thailand |
Antennae | 5-segmented | 5-segmented | 6-segmented | 7-segmented |
Antenna with segment III to remaining segments | shorter | longer | shorter | shorter |
Number of bristles on mandibles | 3 | 4–6 | 3 | 3 |
Number of spines on foretibiae | 4–6 | 6–8 | 8 | 9 |
Carapace flange width | wide | narrow | narrow | very narrow |
Carapace ridges | no | no | no | 2 ridges on each side |
Distal end of carapace | incised notch | incised notch | incised notch | protruding notch |
Length of inner canine to outer canine | shorter than outer canine | shorter than outer canine | subequal | subequal |
Ratio length of carapace (along median suture):width | 0.95–1.00 (n = 3) | 0.96–1.02 (n = 3) | unknown | 1.07–1.13 (n = 3) |
Convexity: carapace depth:length | 0.39–0.45 (n = 3) | 0.30–0.49 (n = 3) | unknown | 0.56–0.58 (n = 3) |
(in alcohol). Body length 1.94–2.43 mm excluding caudal filaments.
Head. Head yellowish with median blackish ocellus between antennae (Fig.
Prosopistoma carinatum sp. n.: A Antenna B–C Labium B ventral view C ventral view D Labrum, dorsal view E–G Mandible E Left mandible (ventral view) F Right mandible (ventral view) G Enlargement of canine of the right mandible, (H). Scale bars: 0.02 mm (H); 0.05 mm (G); 0.1 mm (A, D, E, F); 0.2 mm (B, C).
Carapace. Carapace general colouration dark brown, distally with an irregular (zig-zag) yellow pattern at 0.7 times the length of the carapace (Fig.
Legs. Dorsal margin of fore femur with 14 short simple setae (Fig.
Abdomen. Abdominal gills (Fig.
Unknown.
The name carinatum (Latin for carinate or keeled), refers to the prominent keels or ridge-like mesonotal convexity of the species.
Consensus phylogenetic tree based on the maximum likelihood (ML) analysis of three Thai Prosopistoma species (the best model: T92 +G, parameter = 0.2619). The bootstrap consensus tree inferred from 1,000 replicates. Values above the branches are ML bootstrap values (> 70%). Baetisca laurentina from GenBank was used as the outgroup. There were a total of 658 positions in the final dataset. Abbreviations are the same as those found in Table
The partial sequences of the mitochondrial COI gene (658 bp) of three species found in Thailand were analysed to investigate species delineation. Ten specimens of Prosopistoma were examined. In this study, we used Baetisca laurentina from GenBank (HM902945 and KR144660) as the outgroup since Baetiscidae is the sister family of Prosopistomatidae (Ogden et al. 2009). The consensus phylogenetic tree of ML analysis is shown in Figure
The molecular analysis revealed that Prosopistoma carinatum sp. n. is clearly separated from the other species. The molecular analysis suggests the morphological distinction between the three species. Morphologically, the new species differs from the two previously known species (P. annamense and P. sinense) by (i) longitudinal line ridges on the carapace, (ii) a strongly convex carapace, (iii) the sternal plate, (iv) the scale-like structures on the carapace, (v) 7-segmented antennae and (vi) nine pectinate setae on the ventral margin of the fore-tibiae. The larvae of P. annamense and P. sinense were placed in the same clade, and they share the characteristics of 5-segmented antenna and a smooth carapace. Moreover, the sternal plate of P. annamense and P. sinense has smooth surface (Figs
A comparison of the larvae of all Thai species is shown in Table
The beetle-like mayfly larvae Prosopistoma inhabit shallow water under small stones in streams with a moderate to rapid current (
In Thailand, the Isthmus of the Kra region is a widely recognised as a biogeographic boundary (
This research has been supported by the Centre of Excellence on Biodiversity (BDC) Office of Higher Education Commission (BDC-PG2-161004). We are most grateful to our colleagues for assistance during field trips. The authors would like thank the subject editor, Dr Helen Barber-James, and Dr Frederico Salles for their comments on the manuscript. We would also 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.