Research Article |
Corresponding author: Santi Watiroyram ( santi.watiroyram@npu.ac.th ) Academic editor: Sanda Iepure
© 2021 Santi Watiroyram.
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:
Watiroyram S (2021) Attheyella (Canthosella) thailandica sp. nov. (Copepoda, Harpacticoida, Canthocamptidae) from caves in Thailand. Subterranean Biology 37: 57-73. https://doi.org/10.3897/subtbiol.37.55376
|
During this sampling campaign, the canthocamptid Attheyella (Canthosella) thailandica sp. nov. was collected from various caves in Thailand. The new species is widely distributed in the country and favours habitats, such as phytotelmata and wet soil. Attheyella (Canthosella) thailandica sp. nov. is the second member of the genus to be found in Thailand, after Attheyella (Canthosella) vietnamica
Morphology, phytotelmata, southeast Asia, taxonomy, wet soil
The genus Attheyella Brady, 1880 is found in a wide range of habitats in various water bodies and often in groundwater (
This research mainly focused on cave-dwelling copepods in freshwater from the epikarst zone and related habitats – especially water dripping from rocks and plants at cave entrances (Fig.
Distribution of Attheyella (Canthosella) thailandica sp. nov. A black circle = capital city, Arabic numbers refer to sampling caves: 1, Huang Po cave; 2, Khao Chakan cave; 3, Plub Pleung Thong cave; 4, Payanakarat cave; 5, Pra Hor cave; 6, Mae-nang Songsri cave B Huang Po cave C Plub Pleung Thong cave (white arrow indicates water container with the new species) D Payanakarat cave E Mae-nang Songsri cave F Khao Chakan cave G Pra Hor cave.
All appendages and body ornamentation were examined with 1000× magnification under an Olympus CX31 compound microscope. Drawings were made using an Olympus U-DA drawing tube mounted on the microscope. Final versions of the drawings were done using the CorelDRAW 12.0 graphic programme.
Specimens for scanning electron microscopy (SEM) were dehydrated in an ethanol series (50%, 70%, 80%, 90%, 95%, 100% and 100%) for 15 min at each concentration. Specimens were dried in a critical point dryer and mounted on stubs. Mounted specimens were coated with gold in a sputter-coater. SEM photographs were carried out using a LEO 1450VP scanning electron microscope.
Abbreviations used are: Enp, endopod; Exp, exopod; Exp/Enp-, exopodal segment n/endopodal segment n; P1–P6, legs 1–6; s, spine; a, aesthetasc;
The descriptive terminology follows
Genus Attheyella Brady, 1880
Mae-nang Songsri cave, Hin Tok Subdistrict, Ron Phibun District, Nakhon Si Thammarat Province, southern Thailand: 08°14'45"N, 99°52'01"E, 45 m altitude, 29 October 2015; Payanakarat cave, Tham Thong Lang Subdistrict, Thap Put District, Phang Nga Province, southern Thailand: 08°31'11"N, 98°33'57"E, 140 m altitude, 5 November 2014; Khao Chakan cave, Khao Chakan Subdistrict, Khao Chakan District, Sa Kaeo Province, eastern Thailand: 13°39'36"N, 102°05'04"E, 120 m altitude, 1 September 2017; Plub Pleung Thong cave, Wang Mai Subdistrict, Wang Sombun District, Sa Kaeo Province, eastern Thailand: 13°26'50"N, 102°13'03"E, 223 m altitude, 31 August 2017. All samples were collected by the author.
Huang Po cave, Thung Na Lao Subdistrict, Khon San District, Chaiyaphum Province, north-eastern Thailand: 16°35'25"N, 101°49'28"E, 384 m altitude, 16 October 2017; Pra Hor cave, Tham Yai Subdistrict, Thung Song District, Nakhon Si Thammarat Province, southern Thailand: 08°06'49"N, 99°43'59"E, 101 m altitude, 29 October 2015. Samples were collected by the author.
The specific name of the new species, ‘thailandica’, refers to Thailand, where the species was collected.
Holotype
: one adult female dissected and mounted on one slide,
(holotype). Body (Fig.
Attheyella (Canthosella) thailandica sp. nov., female (A–D) and male (E–G): A habitus, dorsal view B urosome without urosomite 1, dorsal view C urosomite 3–5, ventral view D caudal ramus, dorsal view E cephalothorax, dorsal view F urosome without urosomite 1, dorsal view G urosome without urosomite 1, ventral view.
Caudal ramus
(Figs
Antennule
(Fig.
Antenna
(Fig.
Mandible
(Fig.
Maxillule
(Fig.
Maxilla
(Fig.
Maxilliped
(Fig.
P1–P4 with three-segmented Exp and two-segmented Enp. The armature formula is as follows (Arabic and Roman numerals indicate number of setae and spines, respectively; not including spinules):
Coxa | Basis | Exp | Enp | ||||
1 | 2 | 3 | 1 | 2 | |||
P1 | 0-0 | I-1 | I-0 | I-0 | I-2-1 | 0-1 | 0-2-1 |
P2 | 0-0 | I-0 | I-0 | I-1 | II-2-1 | 0-0 | 0-I+1-0 |
P3 | 0-0 | 1-0 | I-0 | I-1 | II-2-2 | 0-0 | 0-I+2-0* |
P4 | 0-0 | 1-0 | I-0 | I-1 | II-2-2 | 0-0 | 0-2-0 |
P1 (Fig.
P2 (Fig.
P3 (Fig.
P4 (Fig.
P5 (Fig.
P6 (Fig.
Adult females with single egg sac containing 12–15 eggs (holotype: 12 eggs).
(allotype). Body length 510 µm (Fig.
Antennule
(Figs
P1, P4 (Figs
P5 (Figs
(a) The free distal margin of the anal operculum varies from six to ten spinules in females and six to eight spinules in males, a characteristic which is, perhaps, useless for differentiating amongst species, as mentioned by
Attheyella (C.) thailandica sp. nov. is usually found in pools of water at cave entrances, where water seeps through soil and plants before flowing down into the cave (see Fig.
Attheyella (C.) thailandica sp. nov. has been identified as belonging to the subgenus Canthosella Chappuis, 1931 because it shares the following characteristics with other members of the subgenus: posterior margin of somites smooth; caudal ramus longer than wide; antenna with one-segmented Enp; P1–P4 with two-segmented Enp (P1 Enp shorter than Exp, P4 Enp-1 much smaller than other legs of Enp-1); P2–P4 Enp-2 with at least two elements; male P3 with three-segmented Enp bearing an inner apophysis on Enp-2; male P4 Exp-3 without transformed spine; P2–P4 Exp-2–3 with at least one inner seta and one outer spine; female P5 baseoendopod well-developed (reaching beyond Exp) bearing six setae, while Exp bears five setae; male P5 with two setae on baseoendopod and four setae on Exp.
However, A. (C.) thailandica sp. nov. shows morphological differences from other species of the subgenus, related to the ornamentation of the caudal rami. Most species of Attheyella (Canthosella), except A. (C.) antillica and A. (C.) mervini, carry spinules on the inner margin of the caudal ramus (at least in the female). This margin is bare in both sexes of A. (C.) thailandica sp. nov.
At present, the Canthosella subgenus shows two lineages, which include ten American and six SEA species. They can easily be differentiated by the number of armatures on their P2–P4 legs. In contrast to SEA species, all American species, except A. (C.) acanthophora, show a higher number of setae on the P2–P4 Enp-2 in both sexes. Females of these American species have three to four, three to five or two elements on P2–P4, respectively – except for the P4 of A. (C.) antillica, which has one seta. Females of SEA species show one to two setae, one to three setae or one seta on P2–P4, respectively – except for the P4 of A. (C.) vietnamica and A. (C.) thailandica sp. nov., which carry two setae. Additionally, the male P3 Enp-3 of the American species shows two apical setae – except in A. (C.) aliena, which has only one seta. Males of the SEA species have only one seta on the male P3 Enp-3, except for A. (C.) thailandica sp. nov., which has two setae. However, a minute seta located close to a long distal seta on the male P3 Enp-3 could easily have been overlooked in previous descriptions (P.H.C. Corgosinho, personal communication).
Amongst the six SEA species, A. (C.) thailandica sp. nov. is most similar to A. (C.) vietnamica for the following reasons (Table
Morphological differences of the new species and the SEA species of the subgenus Canthosella Chappuis, 1931.
Species Characteristics | A. (C.) fluviatilis | A. (C.) lacustris | A. (C.) muscicola | A. (C.) silvicola | A. (C.) vietnamica | A. (C.) thailandica sp. nov. |
---|---|---|---|---|---|---|
Female | ||||||
Anal operculum | With 8 spinules | With 7 spinules | With 7–10 spinules | With 5–8 spinules | With 6 spinules | With 6–10 spinules |
Caudal ramus (CU) | ||||||
– shape | Conical | Sub-rectangular | Oval | Conical | Conical | Conical |
– Length/wide ratio | < 1.3 | 1.5 | > 1.7 | 1.5 | 1.5 | > 1.7 |
– Inner median margin | With spinules | With spinules | With spinules | With spinules | With spinules | Without spinules |
– Insertion point of seta VII | ?? | ?? | ?? | 3/4 of CU | Distal end of CU | 3/4 of CU |
P2–P4 | ||||||
– segmentation | 2.2.2 | 2.2.2 | 2.2.2 | 2.2.2 | 2.2.1 | 2.2.2 |
– Armature on distal Enp | 1.1.1 | 2.21.1 | 2.22.1 | 2.2.1 | 2.3.2 | 2.3.2 |
P5 | ||||||
– Baseoendopod | With spinules With 1 apical seta | With spinules With 1 apical seta | Without spinules With 1 apical seta | Without spinules With 1 apical seta | Without spinules With 2 apical seta | Without spinules With 1 apical seta |
Male | ||||||
– Armature on P3 Enp-3 | 1 | unknown | 1 | 1 | 1 | 2 |
– Armature on P2 and P4 distal Enp | 13.1 | unknown | 1.1 | 1.24 | 1.2 | 1.2 |
Nevertheless, A. (C.) thailandica sp. nov. also shows strong morphological differences from A. (C.) vietnamica in the following aspects. (a) The P4 Enp is two-segmented in both sexes of A. (C.) thailandica sp. nov. (Fig.
In the female of A. (C.) thailandica sp. nov., the innermost seta of the Exp is located at the (sub)distal margin, while it is obviously located at the inner margin in A. (C.) vietnamica. The new species has a third inner seta located at the apex of the baseoendopod, while this third inner seta and the third outer seta are located apically in A. (C.) vietnamica. The male of A. (C.) thailandica sp. nov. has a more developed P5 baseoendopod, in contrast to A. (C.) vietnamica, whose P5 baseoendopod reaches to one-third of the Exp. Thus, A. (C.) thailandica sp. nov. can be established as its own taxonomical unit new to science.
(female unknown for A. (C.) siolii and A. (C.) striblingi)
1 | P4 Enp one-segmented | 2 |
– | P4 Enp two-segmented | 4 |
2 | Caudal ramus with inner margin produced into curved process | A. (C.) acanthophora |
– | Caudal ramus with normal inner margin | 3 |
3 | P2 Enp-2 with two seta and spine | A. (C.) vietnamica |
– | P2 Enp-2 with four setae and spine | A. (C.) kalima |
4 | P4 Enp-2 with one seta | 5 |
– | P4 Enp-2 with 2–3 setae | 9 |
5 | Caudal ramus with spinules along dorso-inner margin | 6 |
– | Caudal ramus without spinules along dorso-inner margin | A. (C.) antillica |
6 | P2 Enp-2 with one seta | A. (C.) fluviatilis |
– | P2 Enp-2 with two seta and spine | 7 |
7 | P3 Enp-2 with one apical seta and one outer seta | A. (C.) muscicola |
– | P3 Enp-2 with two apical setae | 8 |
8 | P5 baseoendopod with spinules along inner margin | A. (C.) lacustris |
– | P5 baseoendopod without spinules along inner margin | A. (C.) silvicola |
9 | P4 Enp-2 with two seta and spine | 10 |
– | P4 Enp-2 with three setae and spine | A. (C.) pilagaensis |
10 | P2 Enp-2 with two seta and spine | A. (C.) thailandica sp. nov. |
– | P2 Enp-2 with four setae and spine | 11 |
11 | P3 Enp-2 with four setae and spine | 12 |
– | P3 Enp-2 with five setae and spine | 13 |
12 | Caudal ramus with spinules along dorso-inner margin | A. (C.) vera |
– | Caudal ramus without spinules along dorso-inner margin | A. (C.) mervini |
13 | Caudal ramus with dorsal seta at 1/2 of length | A. (C.) chocoensis |
– | Caudal ramus with dorsal seta at 3/4 of length | A. (C.) aliena |
(male unknown for A. (C.) kalima, A. (C.) lacustris and A. (C.) pilagaensis)
1 | P5 baseoendopod with two setae | 2 |
– | P5 baseoendopod unarmed | 7 |
2 | Caudal ramus with inner margin produced into curved process | A. (C.) acanthophora |
– | Caudal ramus with normal inner margin | 3 |
3 | P4 Enp with one apical seta | 4 |
– | P4 Enp with two apical seta and spine | 5 |
4 | P2 Enp-2 with one seta | A. (C.) muscicola |
– | P2 Enp-2 with three setae and spine | A. (C.) fluviatilis |
5 | P3 Enp-3 with one seta | 6 |
– | P3 Enp-3 with two setae | A. (C.) thailandica sp. nov. |
6 | P4 Enp with two subequal apical setae | A. (C.) silvicola |
– | P4 Enp with outer apical seta longer than 2.0× that of inner apical seta | A. (C.) vietnamica |
7 | P4 Enp with one apical seta | A. (C.) antillica |
– | P4 Enp with two apical seta and spine | 8 |
8 | P2 Enp-2 with three setae and spine | 9 |
– | P2 Enp-2 with four setae and spine | 10 |
9 | P3 with two-segmented Enp | A. (C.) mervini |
– | P3 with three-segmented Enp | A. (C.) siolii |
10 | Caudal ramus with spinules along dorso-inner margin | 11 |
– | Caudal ramus without spinules along dorso-inner margin | 12 |
11 | P3 Enp-3 with one apical seta | A. (C.) aliena |
– | P3 Enp-3 with two apical setae | A. (C.) vera |
12 | Caudal ramus slightly longer than wide | A. (C.) chocoensis |
– | Caudal ramus about 2.0× longer than wide | A. (C.) striblingi |
The author would like to thank the National Research Council of Thailand (Grant No. 2559A13402007; 2560A13402010; 256108A1340006) for financial support. The author would like to thank the inputs of Dr Santiago Gaviria (Vienna) and Dr Paulo Henrique Costa Corgosinho for suggestions to correct the manuscript.