Research Article |
Corresponding author: Supiyanit Maiphae ( supiyanit.m@ku.ac.th ) Academic editor: Kay Van Damme
© 2023 Thanida Saetang, Supiyanit Maiphae.
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:
Saetang T, Maiphae S (2023) Diversity of the genus Tropodiaptomus Kiefer, 1932 (Crustacea, Copepoda, Calanoida, Diaptomidae) in Thailand, with the description of two new species. Zoosystematics and Evolution 99(2): 399-422. https://doi.org/10.3897/zse.99.105511
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Tropodiaptomus is a genus of diaptomid copepods with 10 species currently recorded in Thailand. A recent study on DNA taxonomy revealed putative new species among specimens collected from freshwater habitats throughout Thailand. This study examined the morphological characteristics and confirmed the taxonomic status of the two putative new species of Tropodiaptomus. Results showed that the two new taxa were different from other species in the genus Tropodiaptomus. These two new species, T. pedecrassum sp. nov. and T. longiprocessus sp. nov., were described and illustrated based on material collected from a swamp in northern Thailand and a pond in western Thailand, respectively. They were distinguished from their congeners by the length of the spinous process on the antepenultimate segment of the adult male right antennule, the number of lobes and serration pattern on the inner margin of the adult male left P5, and the shape and supplementary process on the surface structures of basis and distal exopod segments of the adult male right P5. These discoveries increased the number of records of this genus in Thailand to 12 species. A pictorial key to all species is provided, and their ecological and biogeographical distributions are updated and discussed.
habitat preference, new species, Thailand, Tropodiaptomus longiprocessus, Tropodiaptomus pedecrassum
To date, 170 copepod species have been recorded from freshwater habitats in Thailand (
Our study proposes Tropodiaptomus sp. 1 and Tropodiaptomus sp. 2 mentioned in
Samples were qualitatively collected from swamp in Dok Kham Tai District, Phayao Province, northern Thailand (19°13'57.6"N, 100°02'56.5"E) in January 2018 and Thong Phaphum District, Kanchanaburi Province (14°39'09.1"N, 98°33'27.5"E) in June 2019 using a plankton net of 60 µm mesh and immediately preserved in 70% ethanol. All adult males and females were sorted using an Olympus SZ40 stereo microscope, and each specimen was dissected and mounted on a slide in glycerine and then sealed using nail varnish.
The morphological characteristics were examined and identified using an Olympus CH2 compound microscope, and drawings were made from both complete and dissected specimens using a camera lucida connected to the microscope. The final versions of the drawings were made using Adobe Illustrator CS5 program (version 15.0). The specimens were identified to species level according to
All type specimens were deposited in the reference collection of the Princess Maha Chakri Sirindhorn National History Museum, Prince of Songkla University, Songkhla, Thailand (PSUNHM).
Order CALANOIDA Sars, 1903
Family DIAPTOMIDAE Baird, 1850
Tropodiaptomus orientalis (Brady, 1886).
Swamp near rice field, Dok Kham Tai District, Phayao Province, northern Thailand (19°13'57.6"N, 100°02'56.5"E). Temporary habitat without macrophytes.
Holotype. One adult male, dissected and mounted onto five slides, Dok Kham Tai, Phayao province, northern Thailand (19°13'57.6"N, 100°02'56.5"E), 31 January 2018, Thanida Saetang and Supiyanit Maiphae; PSUZC-PK2008-01–PSUZC-PK2008-05. Allotype. One adult female, collected with holotype; PSUZC-PK2008-06–PSUZC-PK2008-08. Paratype. One adult male, collected with holotype; PSUZC-PK2008-09–PSUZC-PK2008-11.
Body
(Figs
Tropodiaptomus pedecrassum sp. nov., male holotype. A. Habitus, dorsal view; B. Segment 1–15 of left antennule (gray color indicates spine); C. Segment 16–25 of left antennule; D. Segment 12–14 of left antennule of the adult male specimen from the same population of holotype, (arrowhead indicates variation of setae on segment 13); female allotype; E. Habitus, dorsal view. Scale bars: 100 µm.
Tropodiaptomus pedecrassum sp. nov., male holotype. A. Pediger 5 and urosome, dorsal view; female allotype; B. Pediger 5 and urosome, dorsal view (arrowhead indicates anterior part swollen in both sides and triangular-like lobe, and grey ellipse indicates dorsal prominence). Scale bar: 100 µm.
Rostrum
(Fig.
Tropodiaptomus pedecrassum sp. nov., male holotype. A. Rostrum; B. Segment 1–11 of right antennule; C. Segment 12–19 of right antennule (gray color indicate spine); D. Segment 20–22 of right antennule; E. Segment 13 of right antennule of the adult male specimen from the same population of holotype (arrowhead indicates hyaline knob). Scale bars: 100 µm.
A1
(Figs
A2
(Fig.
Mandible
(Fig.
Maxillule
(Fig.
Coxal endite with three plumose setae, and coxal epipodite with seven plumose setae and two bipinnate spines. Basis with two endites; the proximal with four plumose setae and the distal with eight plumose setae, and basal exite with one bipinnate spines. Exopod 1-segmented with six plumose setae, one longitudinal row of setules on inner margin of segment. Endopod 1-segmented with four plumose setae.
Maxilla
(Fig.
Maxilliped
(Fig.
P1–P4
(Fig.
Armature formula of P1–P4 in T. pedecrassum sp. nov. and T. longiprocessus sp. nov. (Arabic numerals representing setae and Roman numerals representing spine from outer-inner or outer-apical-inner margins).
Swimming legs | Coxa | Basis | Exp | Enp | ||||
---|---|---|---|---|---|---|---|---|
1 | 2 | 3 | 1 | 2 | 3 | |||
P1 | 0–1 | 0–0 | I–1 | 0–1 | I,3,2 | 0–1 | 1,2,3 | – |
P2 | 0–1 | 0–0 | I–1 | I–1 | I,3,3 | 0–1 | 0–2 | 2,2,3 |
P3 | 0–1 | 0–0 | I–1 | I–1 | I,3,3 | 0–1 | 0–2 | 2,2,3 |
P4 | 0–1 | 1–0 | I–1 | I–1 | I,3,3 | 0–1 | 0–2 | 2,2,3 |
P5
(Figs
Body
(Figs
A1, A2, mandible, maxillule, maxilla, maxilliped, P1–P4 and rostrum (not shown) same as male.
P5
(Figs
Morphological variability has been observed in: (i) the total body length (except of caudal setae) which ranged from 1,145–1,380 µm (mean 1,283 µm, n = 6) in the adult males and 1,520–1,626 µm (mean 1,575 µm, n = 5) in the adult females; (ii) the length of the spinous process on antepenultimate segment of the adult male right A1 is 3/4 to equal of segment 21 (Fig.
Comparison of four morphologically similar species of Tropodiaptomus (after
Morphological characters | T. pedecrassum sp. nov. | T. longiprocessus sp. nov. | T. hebereri | T. lanaonus |
---|---|---|---|---|
The adult male | ||||
Body length, except caudal satae (μm) | 1,145–1,380 | 1,490–1,545 | 1,350–1,400 | 1,120–1,150 |
Right A1: segment-16 with spinous process | Yes | Yes | No | No |
Right A1: relative length of the spinous process of segment 20 | 3/4 of or equal to segment 21 | Longer than segment 21 | 1/2 of segment 21 | Longer than segment 21 |
Left A1: number of setae on segment-13 | 1 or 2 | 1 | 1 | 1 |
Right P5: basis, supplementary process and surface structures | One round process, one semicircular process and one longitudinal hyaline lamella | One triangular process and one longitudinal hyaline lamella | One triangular process, one semicircular process and one semicircular hyaline lamella | One triangular process and one longitudinal hyaline lamella |
Right P5: exp-1, length of spinous process on outer distal margin/ segment length | Equal | Shorter | Shorter | Shorter |
Right P5: exp-2, shape | Rhomboid | Rectangular | Trapezoidal | Rectangular |
Right P5: exp-2, supplementary process | Triangular-shape process, large | Large and long process | Triangular process | Semicircular process |
Right P5: lateral spine of the exp-2 | Short | Long | Short | Long |
Right P5: exp-2, one round hyaline prominence inserted between end claw and lateral spine | Present | Absent | Present | Absent |
Right P5: length of enp vs exp-1 length | Shorter | Longer | Longer | Longer |
Left P5: exp, inner margin | 1 lobe, the serration gradually decreasing in size from proximal to distal end | 2 lobes, shallow, uniform serration | 1 lobe, the serrate of the proximal part larger than the distal part | 2 lobes, shallow, uniform serration |
Left P5: enp (number of segment) | 2 (yet incompletely separated) | 2 | 1 | 2 |
Left P5: length of finger vs thumb length | Equal | Equal | 3/4 | Equal |
The adult female | ||||
Body length, except caudal satae (μm) | 1,520–1,626 | 1,570–1,700 | 1,500–1,600 | 1,200–1,250 |
Fifth pediger: lateral wings | Asymmetrical | Asymmetrical | Asymmetrical | Symmetrical |
Fifth pediger: dorsal prominence | Present | Absent | Present | Absent |
Genital double-somite | Asymmetrical, dilate on proximal both sides | Asymmetrical, dilate on proximal right side | Asymmetrical, dilate on proximal both sides | Symmetrical |
Genital double-somite: outer distal corner with lobe | Triangular lobe | Absent | Large round lobe | Small round lobe |
Caudal rami: outer margin with setules | Present | Present | Absent | Present |
P5: exp-3 fused with exp-2 | Yes | Yes | No | No data |
P5: end claw, outer margin with setules | Present | Present | Present | No data |
P5: enp, elements on distal end | Spiniform smooth setae | Spiniform smooth setae | Slender smooth seta | Slender smooth seta |
Tropodiaptomus pedecrassum sp. nov., male. A, B. P5, dorsal view; c, d. Exopodal and endopodal segment of left P5; female. E. Pediger 5 and urosome, lateral view; F. Pediger 5 and urosome, dorsal view; g. P5, dorsal view; h. Endopod of P5; i. Exp-2 and exp-3 of P5. Arrowheads show structures discussed in the description.
The specific name ‘pedecrassum’ is derived from the chubby shape of the adult male P5 that is clearly different from the more rectangular shape in other species of the genus.
In our samples, the new taxon co-occurred with one other copepod species, Mongolodiaptomus botulifer (Kiefer, 1974).
Tropodiaptomus pedecrassum sp. nov. was found only in its type locality so far. It was recorded in two out of 471 samples collected from 206 freshwater habitats throughout Thailand between September 2017 and July 2019. Water temperature 19.6 °C, conductivity 620 µs cm-1, salinity 0.3 ppt, total dissolved solids 450 mg L-1, dissolved oxygen 3.3 mg L-1, pH 7.1, and water depth 30–40 cm, substrate with mud.
Tropodiaptomus pedecrassum sp. nov. is confirmed to belong to the genus Tropodiaptomus based on the combination of characteristics mentioned by
This species differs from the congeneric species by the following characters: (i) antepenultimate segment of the male right antennule with straight spinous process reaching 3/4 or equal of next segment; (ii) inner margin of exopodal segment of the male left P5 with single lobe and the serration gradually decreases in size from the proximal to distal end; (iii) basis of the male right P5 with two processes and one longitudinal hyaline lamella; (iv) exp-1 of the male right P5 with triangular lobe on inner margin, distal outer corner produced into acute spinous process, length about as long as its segment; and (v) exp-2 of the male right P5 with rhomboid shape, dorsal surface with one semicircular hyaline knob on proximal outer margin, one semicircular hyaline lamella on distal inner margin, and one triangular process in middle of segment.
According to the identification key given by
Nong Ping swamp (NP2), Thong Phaphum district, Kanchanaburi province, western Thailand (14°38'49.1"N, 98°33'48.8"E). Temporary habitat without macrophytes.
Nong Ping swamp (NP3) Thong Phaphum District, Kanchanaburi Province, Western Thailand (14°39'00.4"N, 98°34'33.7"E). Temporary habitat without macrophytes.
Holotype. Adult male, dissected and mounted onto one slide, Thong Phaphum District, Kanchanaburi province, western Thailand (14°38'49.1"N, 98°33'48.8"E), 22 June 2019, Thanida Saetang and Supiyanit Maiphae; PSUZC-PK2009-01. Allotype. One adult female, collected with holotype; PSUZC-PK2009-02. Paratype. One adult male, collected with holotype; PSUZC-PK2009-03.
Body
(Fig.
Rostrum
(Fig.
A1
(Figs
Tropodiaptomus longiprocessus sp. nov., male holotype. A. Segment 1–11 of right antennule; B. Segment 12–19 of right antennule (gray color indicates spine); C. Segment 20–22 of right antennule; D. Segment 20 and 21 of right antennule of the adult male specimen from the same population of holotype. Scale bar: 100 µm.
A2
(Fig.
Mandible
(Fig.
Maxillule
(Fig.
Maxilla
(Fig.
Maxilliped
(Fig.
P1–P4
(Figs
P5
(Figs
Tropodiaptomus longiprocessus sp. nov., male holotype. A. P5, dorsal view; B–D. Shape variation of hyaline lamella on exp-2 of right P5 in adult male, dorsal surface; E. Exp-2 of right P5 of the adult male specimen from the same population of holotype, lateral view (arrowhead indicates hyaline lamella); Tropodiaptomus longiprocessus sp. nov., female allotype: F. P5 (arrowhead indicates exp-3). Scale bar: 100 µm.
Body
(Figs
Tropodiaptomus longiprocessussp. nov; male. A. P5, dorsal view; B, C. Hyaline lamella on exp-2 of right P5, dorsal surface; D. Hyaline lamella on exp-2 of right P5, lateral surface; E, F. Exopodal segment of the male left antennule,dorsal surface; G. Exopodal segment of the male left antennule,ventral surface; H. Antepenultimate segment of right antennule; I. Lateral part of P1 basis; female: J. Pediger 5 and urosome, lateral view; K. Pediger 5 and urosome, dorsal view; L. P5. Arrowheads show the characters discussed in the description.
A1, A2, mandible, maxillule, maxilla, maxilliped, P1–P4 (not shown) and rostrum (Fig.
P5
(Figs
Morphological variability has been observed in: (i) the total body length (except of caudal setae) which ranged from 1,490 –1,545 µm (mean 1,510 µm, n = 4) in the adult males and 1,570–1,700 µm (mean 1,646 µm, n = 6) in the adult females; and (ii) shape and length of large and long process on exp-2 of the adult male right P5 (Figs
The specific name ‘longiprocessus’ is derived from the presence of a long process on exp-2 of the right P5 in the adult male.
There were no other diaptomid copepods in these samples.
Tropodiaptomus longiprocessus sp. nov. was found only in its type locality. It is rare because it was found only in two of 471 samples collected from 206 freshwater habitats throughout Thailand between September 2017 and July 2019.Water temperature 26.6–28.0 °C, conductivity 623.3–672.7 µs cm-1, salinity 0.3 ppt, total dissolved solids 385–424 mg L-1, dissolved oxygen 2.5–6.0 mg L-1, pH 7.2–7.3, and water depth 0.2–0.3 m, substrate with mud. However, in order to understand more in their habitat preference, the whole year samples are needed to be examined.
Tropodiaptomus longiprocessus sp. nov. differs from the congeneric species by the following respects: (i) antepenultimate segment of the male right antennule with straight spinous process reaching beyond the distal margin of next segment; (ii) inner margin of exopodal segment of the male left P5 with two lobes, and with uniform serration; (iii) basis of the male right P5 with one triangular process and one longitudinal hyaline lamella; (iv) exp-1 of the male right P5 with triangular lobe on inner margin, distal outer corner produced into acute spinous process; and (v) exp-2 of the male right P5 rectangular, dorsal surface with two longitudinal hyaline lamellae in middle and distal inner margin and one large and long process in middle in lateral third.
Tropodiaptomus longiprocessus sp. nov. is the most similar to T. lanaonus, but it differs distinctively in the following characters: (i) the dorsal surface of exp-2 of the adult male right P5 has two longitudinal hyaline lamellae and one large and long process; (ii) lateral wings of the fifth pediger of adult female are asymmetrical; (iii) anterior part of the genital double-somite of the adult female is dilated on the right side; (iv) right posterior corner of the genital double-somite of the adult female does not have a round lobe; and (v) distal end of the endopodal segment of the adult female P5 has smooth spiniform setae (Table
The diversity of Tropodiaptomus in Thailand is presented as a pictorial key in Fig.
Tropodiaptomus species have been recorded from various ponds, lakes, streams, roadside canals and rice field habitats in Thailand (
Geographic distribution of Tropodiaptomus species in Thai water bodies and remarks on their status.
Species | Distribution in Thai water bodies | Co-occurring species in Thai water bodies | Wider Distribution | Notes |
---|---|---|---|---|
1. T. doriai | Freshwater habitat close to Bangkok, central Thailand | No data | India, Indonesia and Sri Lanka ( |
It was recorded only once in Thailand ( |
2. T. hebereri | Cold season (January), in a roadside canal in northern Thailand | None | China, India, Indonesia and Malaysia ( |
|
3. T. lanaonus | Floodplain, lake and pond in both dry (March) and rainy seasons (August) in central and northeastern Thailand | No data | Philippines ( |
|
4. T. cf. lanaonus | Rice field, river, roadside canal, swamp in both dry (May) and rainy seasons (June, September, October, November) in eastern, northeastern and southern Thailand | Dentodiaptomus javanus, Eodiaptomus phuphanensis, Heliodiaptomus elegans, Mongolodiaptomus botulifer, M. dumonti, M. malaindosinensis, M. pectinidactylus, Phyllodiaptomus christineae, P. roietensis, P. surinensis, Phyllodiaptomus sp., Tropodiaptomus megahyaline, T. vicinus, Vietodiaptomus blachei | – | This species does not agree with the original description of T. lanaonus in (i) the length of the spinous process in the antepenultimate segment of adult male right antennule is 1/2 to 3/4 of segment 21, and (ii) the ornamentation on the basis of adult male right P5 has one hyaline lamella and one apophysis or one hyaline lamella and no apophysis ( |
5. T. megahyaline | Pond and rice field in both dry (May) and rainy season (June) in northeastern Thailand | Heliodiaptomus elegans, Mongolodiaptomus malaindosinensis, M. pectinidactylus, Neodiaptomus songkhramensis, Tropodiaptomus cf. lanaonus, Vietodiaptomus blachei | Endemic in Thailand ( |
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6. T. oryzanus | Several types of habitats including canal, floodplain, lake, man-made pond, pond, rice field and roadside canal in dry (April, May), rainy (June, August, October) and cold season (December) in northeastern Thailand | Dentodiaptomus javanus, Neodipatomus laii, Mongolodiaptomus botulifer, M. malaindosinensis, Tropodiaptomus vicinus | Cambodia, China, Japan, Korea, Malaysia, Taiwan and Vietnam ( |
|
7. T. ruttneri | Peat swamp, swamp roadside canal and pond in both dry (March) and rainy seasons (August, November) in north, northeastern and southern Thailand | Mongolodiaptomus botulifer | China and Malaysia ( |
|
8. T. vicinus | Several types of habitats including canal, fish pond, floodplain, lake, man-made pond, marsh, peat swamp, pond, rice field, river, swamp and roadside canal in dry (April, May), rainy (June, August, October, November) and cold season (December) in the central, eastern, northeastern, southern and western Thailand | Dentodiaptomus javanus, Eodiaptomus phuphanensis, Heliodiaptomus elegans, Mongolodiaptomus botulifer, M. malaindosinensis, Neodipatomus laii, N. yangtsekiangensis, Phyllodiaptomus christineae, P. roietensis, P. surinensis, Tropodiaptomus oryzanus, T. cf. lanaonus | Cambodia, India, Indonesia, Laos, Malaysia, Philippines and Vietnam ( |
|
9. T. pedecrassum sp. nov. | Swamp in cold season (January, February) in northern Thailand | Mongolodiaptomus botulifer | Recorded as Tropodiaptomus sp.1 in |
|
10. T. longiprocessus sp. nov. | Swamp in rainy season (June) in western Thailand | None | Recorded as Tropodiaptomus sp.2 in |
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11. Tropodiaptomus sp. 1 | Pond in rainy season (October) in northeastern Thailand | None | Recorded as Tropodiaptomus sp.3 in |
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12. Tropodiaptomus sp. 2 | Pond in northern Thailand no information available on temporal distribution | No data | Recorded as Tropodiaptomus sp. in |
Each species of Tropodiaptomus can be found in various types of freshwater habitats, while most occur in both temporary and permanent water bodies with a limited geographical distribution range (Fig.
This study assessed the morphological characteristics and molecular data recorded by
The discovery of these two new species increases the number of Tropodiaptomus species recorded in Thailand from 10 to 12, thus accounting for 19% of species diversity worldwide. Based on the large number of samples taken throughout Thailand, most Tropodiaptomus species have a limited distribution at one site or one region, with only T. vicinus having a wide distribution. Moreover, Tropodiaptomus is distributed in a specific habitat, temporary pools without vegetation and the occurrences of this genus are seasonal. This type of habitat is undersampled in SE Asia, however it is important for our understanding of the diversity of this genus. However, recent findings of new records in Southeast Asia (
We would like to thank the editor and the reviewer for taking the time and effort to review the manuscript. We sincerely appreciate all the valuable comments and suggestions that helped us improve the quality of the manuscript. This research was approved by the Institutional Animal Care and Use Committee, Kasetsart University, Thailand (approval no. ACKU61-SCI-004). The authors would like to thank Kasetsart University for providing post-master research scholarship through the Biodiversity Center Kasetsart University. Dr. Rapeepan Jaturapruek is thankful for assistance in field sampling.