Research Article
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Research Article
Attheyella (Canthosella) thailandica sp. nov. (Copepoda, Harpacticoida, Canthocamptidae) from caves in Thailand
expand article infoSanti Watiroyram
‡ Nakhon Phanom University, Nakhon Phanom, Thailand
Open Access

Abstract

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 Borutzky (1967), which is most similar to it. Amongst Asian species, both A. (C.) thailandica sp. nov. and A. (C.) vietnamica have identical setal formulae, with a greater number of armatures on the distal endopods of legs 2–4. However, A. (C.) thailandica sp. nov. markedly differs from A. (C.) vietnamica in the insertion point of the dorsal seta and the presence of inner spinules on the caudal ramus. Additionally, the leg 4 endopod is two-segmented in A. (C.) thailandica sp. nov., but one-segmented in A. (C.) vietnamica.

Keywords

Morphology, phytotelmata, southeast Asia, taxonomy, wet soil

Introduction

The genus Attheyella Brady, 1880 is found in a wide range of habitats in various water bodies and often in groundwater (Chang and Kim 1992). The subgenus Attheyella (Canthosella) Chappuis, 1931 is one of the five subgenera of Attheyella and contained 15 valid species: A. (C.) acanthophora (Delachaux, 1924); A. (C.) aliena Noodt, 1956; A. (C.) antillica (Petkovski, 1973); A. (C.) chocoensis Gaviria & Defaye, 2012; A. (C.) fluviatilis Chappuis, 1931; A. (C.) kalima (Delachaux, 1924); A. (C.) lacustris Chappuis, 1931; A. (C.) mervini Janetzky, Martinez Arbizu & Reid, 1996; A. (C.) muscicola (Chappuis, 1928); A. (C.) pilagaensis Janetzky, Martinez Arbizu & Reid, 1996; A. (C.) silvicola Löffler, 1973; A. (C.) siolii (Kiefer, 1967); A. (C.) striblingi (Reid, 1990); A. (C.) vera Por & Hadel, 1986 and A. (C.) vietnamica Borutzky, 1967 (Walter and Boxshall 2020). To date, six species of the subgenus Canthosella have been originally described from southeast Asia (SEA): three species from Indonesia [A. (C.) fluviatilis, A. (C.) lacustris and A. (C.) muscicola] and one species each from Malaysia [A. (C.) silvicola], Vietnam [A. (C.) vietnamica] and Thailand [A. (C.) thailandica sp. nov.], respectively. In Thailand, the first Attheyella species found in a cave, A. (C.) vietnamica, has being collected in northern Thailand (Watiroyram et al. 2015) and the second species of the genus found in a cave of the country is described herein as A. (C.) thailandica sp. nov.

Material and methods

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. 1). Samples were collected using a filtering bottle with a mesh size of 60 µm and were preserved immediately in 70% ethanol. In the laboratory, samples were rinsed with tap water through a sieve with 60 µm mesh size. Adult specimens were sorted under an Olympus SZ51 stereomicroscope at 40× magnification and were placed in a mixture of glycerol and 70% ethanol (ratio ~ 1:10 v/v) to pure glycerol. Animals were dissected and prepared on a glycerine-mounted slide under a stereomicroscope at 40–100× magnification. The specimens were mounted in pure glycerine on a glass slide and were sealed under a cover glass with transparent nail varnish. Whole specimens were stored in 70% ethanol.

Figure 1. 

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; P1P6, legs 1–6; s, spine; a, aesthetasc; NHMUK, the Natural History Museum (United Kingdom); NPU, Nakhon Phanom University, Faculty of Science (Thailand).

The descriptive terminology follows Huys and Boxshall (1991).

Taxonomic section

Family Canthocamptidae Brady, 1880

Genus Attheyella Brady, 1880

Subgenus Canthosella Chappuis, 1931

Attheyella (Canthosella) thailandicasp. nov.

Figures 2, 3, 4, 5, 6, 7

Other localities

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.

Type localities

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.

Etymology

The specific name of the new species, ‘thailandica’, refers to Thailand, where the species was collected.

Type specimens

Holotype : one adult female dissected and mounted on one slide, NHMUK 2020.56; Allotype: one adult male dissected and mounted on one slide, NHMUK 2020.57; Paratypes: three adult females and three adult males not dissected and stored in a 1.5 ml microtube with 70% ethanol, NHMUK 2020.58-63; one adult female dissected and mounted on one slide, NPU 2020-003; one adult male dissected and mounted on one slide, NPU 2020-004; three adult females and three adult males not dissected and stored in a 1.5 ml microtube with 70% ethanol, NPU 2020-005.

Description of adult female

(holotype). Body (Fig. 2A) cylindrical, with mean length 510 µm, measured from rostrum to distal rim of caudal rami (n = 5, range 500–530 µm). Cephalothorax with narrow, saddle-shaped, well discernible integumentary window; several sensilla scattered on dorso-lateral surface. Prosome and urosome (segments 1–4) with free posterior margins smooth dorsally; rows of minute spinules dorsally on prosomites 2–4, urosomites 1–2. Genital double-somite completely fused (Figs 2B, 4A), about 0.5× as long as wide, with row of strong dorso-lateral and ventral spinules along distal margin, mid-ventral bell-shaped copulatory pore, duct and receptacles. Urosomite 3 with row of spinules ventrally inserted near posterior edge, row interrupted mid-ventrally. Urosomite 4 with continuous row of spinules ventrally near posterior margin. Anal somite (Figs 2B, C, 4A, B) with one pair of sensilla dorsally above base of anal operculum; seven to ten spinules (nine in holotype) ventrally near insertion of each caudal ramus; anal operculum concave and well-developed, with six strong spinules on free posterior margin.

Figure 2. 

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 2A–D, 4A, B) conical, about 2.0× longer than wide, inner margin unornamented; longitudinal keel located along dorso-inner margin, ending in acute tip. Ramus with seven setae (setae I–VII), all smooth, except setae IV and V. Anterolateral accessory seta (I) small, inserted near seta II. Anterolateral seta (II) inserted at two-quarters of ramus, accompanied by two spinules, about 1.5× longer than ramus length. Posterolateral seta (III) inserted at three-quarters of ramus, accompanied by two to three (two in holotype) spinules, about 1.3× longer than ramus length. Outer apical seta (IV) unipinnate, without a breaking plane, about 1.7× longer than ramus length. Inner apical seta (V) longest, bipinnate, without a breaking plane, more than 5.0× longer than ramus length. Inner accessory seta (VI) shortest, about as long as ramus length. Dorsal seta (VII) articulated, inserted on distal end of dorsal keel at three-quarters length of ramus, about 1.6× longer than ramus length.

Antennule (Fig. 3A) eight-segmented, with setal formulae as follow: 1(I), 8(II), 5(III), 1+a(IV) with conjoined bases, 1(V), 2(VI), 2(VII), 7 and 1+a(VIII) with conjoined bases (Arabic and Roman numerals refer to number of setae and segment number, respectively). Aesthetasc cylindrical; both aesthetascs fused at their bases with a seta forming an acrotheck. All setae slender, smooth.

Figure 3. 

Attheyella (Canthosella) thailandica sp. nov., female: A antennule B antenna C mandible D maxillule E maxilla F maxilliped.

Figure 4. 

Attheyella (Canthosella) thailandica sp. nov., female (A, B) and male (C): A urosome without urosomite 1, ventral view (A' variation on urosomite 3–4, from type locality) B, C anal somite and caudal ramus, lateral view.

Antenna (Fig. 3B) with small, unarmed coxa. Allobasis elongated, with two spinules on abexopodal margin. Exp one-segmented, with two inner and two apical unipinnate setae. Enp one-segmented, with two outer spines, one apical spine and five apical setae (two normal and three geniculated setae); inner and outer margins ornamented with two rows of spinules.

Mandible (Fig. 3C) with two large teeth, seven small teeth distally and one lateral seta on gnathobase, with a small knob on the disto-lateral margin. Palp one-segmented, with four apical setae.

Maxillule (Fig. 3D) with five apical spines and one anterior seta on praecoxal arthrite. Coxal endite with one spine and one seta apically. Basis with one spine, two setae apically. Exp and Enp reduced, represented by four lateral setae on basis.

Maxilla (Fig. 3E) with two endites on syncoxa, each endite with three apical setae. Basis elongated, drawn out into a claw, with one proximal accessory seta. Enp reduced and represented by two setae.

Maxilliped (Fig. 3F) with unarmed coxa, ornamented with row of spinules on median surface. Basis with two groups of spinules at inner and outer margins. Enp one-segmented, transformed into claw-like segment, accompanied by one small seta inserted proximally.

P1P4 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. 5A) basis with short, strong outer spiniform spine and long, slender, smooth inner seta. Enp shorter than Exp; Enp-1 reaching to middle of Exp-2, with one inner unipinnate-tipped seta, ornamented with strong outer spinules and inner setules. Enp-2 with one smooth distal inner seta, one distal inner geniculated seta and one distal outer unipinnate seta; ornamented with strong outer spinules. Exp-1–2 with one distal outer spiniform spine; both segments ornamented with strong outer spinules. Exp-3 with one distal outer spiniform spine, two apical setae (inner seta geniculated, outer one unipinnate) and one distal inner geniculated seta; ornamented with few distal outer spinules.

Figure 5. 

Attheyella (Canthosella) thailandica sp. nov., female: A P1 B P2 C P3 D P4 (D' variation) E P5 (E' variation, specimen from Pra Hor cave).

P2 (Fig. 5B) basis with short, strong outer spine. Enp as long as Exp-1; Enp-1 shorter than wide, unarmed. Enp-2 with two elements apically; outer spine slightly longer than segment; inner seta bipinnate, long, extending beyond Exp. Exp-1 with one enlarged distal outer spine; ornamented with outer spinules. Exp-2 with one distal outer spine and one smooth distal inner seta. Exp-3 with two distal outer spines, two apical setae (inner seta bipinnate, outer one unipinnate) and one smooth inner seta; ornamented with disto-outer spinules.

P3 (Fig. 5C) basis with long, smooth, slender outer seta. Enp as long as Exp-1; Enp-1 shorter than wide, unarmed. Enp-2 with three elements apically; outer spine and innermost seta shorter than segment, subequal in length; middle seta bipinnate, reaching beyond Exp. Exp-1–2 with one distal outer spine, ornamented with spinules along disto-outer margin; Exp-2 with one additional smooth distal inner seta. Exp-3 with two distal outer spines, two apical setae (inner seta bipinnate, outer one unipinnate) and two inner setae (distal seta unipinnate, about 3.0× longer than proximal seta); ornamented with spinules along disto-outer margin.

P4 (Fig. 5D) basis and Exp as in P3, but Exp-3 shorter. Enp as long as Exp-1; Enp-1 small, shorter than wide, unarmed. Enp-2 with two bipinnate setae apically, both longer than segment; inner seta shorter than outer seta.

P5 (Fig. 5E) without ornamentation on surface. Basal seta smooth, slender. Baseoendopod separated from Exp, well-developed, exceeding Exp, with six spiniform setae; third inner seta longest apically; remaining setae decreasing in length to outer and inner margins of Enp. Exp sub-oval, with five setae, second inner seta longest; two innermost setae spiniform; three outer remaining setae smooth and decreasing in length to margin of Exp.

P6 (Fig. 4A) reduced to a single bipinnate seta inserted on the small plate, anterior to the seminal receptacle on the first half of genital double-somite.

Adult females with single egg sac containing 12–15 eggs (holotype: 12 eggs).

Description of adult male

(allotype). Body length 510 µm (Fig. 2E–G) from rostrum to distal rim of caudal ramus, 470–590 µm (n = 5), smaller than female. Prosome (including antenna and mouthparts), anal somite and caudal ramus similar to those of female. Genital somite (Figs 2F, 6B) without row of posterior spinules; urosomites 3–5 with continuous posterior spinules along ventral to dorso-lateral sides.

Antennule (Figs 67A) ten-segmented, geniculated between segments 7 and 8. Segment 4 small, beneath segment 3. Setal formulae: 1(I), 7(II), 5(III), 2(IV), 3 and 1+a(V), 2(VI), 1(VII), 0(VIII), 0(IX), 7 and 1+a(X); aesthetasc on segments 5 and 10 fused to the base of seta, forming an acrotheck. All setae smooth.

Figure 6. 

Attheyella (Canthosella) thailandica sp. nov., male: A antennule (white arrow indicates segment IV) B P4 Enp (white arrow indicates Enp-1) C P5 and P6.

P1, P4 (Figs 6B, 7D) and P2–3 Exp similar to those in female. P2 (Fig. 7B) Enp slightly longer than Exp-1; Enp-1 shorter than wide, unarmed. Enp-2 with long bipinnate seta apically, with two to three spinules on outer margin. P3 (Fig. 7C) Enp three-segmented; Enp-1 shorter than wide, unarmed. Enp-2 with thin inner apophysis with harpoon-like tip, long, exceeding beyond Exp. Enp-3 with two apical setae; outer seta bipinnate, longer than Exp; inner seta smooth, thin, shorter than segment. P4 (Fig. 7D) Enp two-segmented; Enp-1 small, shorter than wide, unarmed; Enp-2 shorter than Exp-1, about 2.5× longer than wide, with two bipinnate setae apically (outer seta longer than inner one; inner seta slightly longer than segment).

Figure 7. 

Attheyella (Canthosella) thailandica sp. nov., male: A antennule B P2 (B' variation, specimen from type locality) C P3 D P4 Enp (D' variation, specimen from Pra Hor cave) E P5.

P5 (Figs 6C, 7E) separated from somite, baseoendopod of left and right sides fused medially. Basal seta long, slender and smooth. Baseoendopod separated from Exp, reaching one-half of Exp, with two spiniform setae; inner seta over 4.0× longer than outer seta. Exp with four setae; second inner seta longest, followed by second outer seta, innermost seta and outermost seta, respectively; two inner setae bipinnate, two outer setae smooth.

Variability

(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 Gaviria and Defaye (2012). (b) The posterior margin of urosomite 3 in female has a continuous row of ventral spinules (Fig. 4A'; one of the other five examined specimens collected from the type locality). (c) The P2P3 Enp-2 has a different number of spinules along the outer margin, with two spinules on P2 and three spinules on P3 in females (not shown in Figure; one of the other five female from the type locality) and with two spinules on P2 in males (Fig. 7B'; one of the other five examined specimens collected from the type locality). (d) The seta size of the P4 Enp-2 in both sexes, the outer seta about 2.0× longer than the inner seta in female and the inner seta longer than or equal in length to outer seta in male (Figs 5D', 7D'; two of the other five examined specimens collected from Pra Hor cave). (e) The female P5 in the southern population shows mostly shorter Exp and End setae (Fig. 5E'; two other females from Pra Hor cave).

Ecology

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. 1B, D, G). In this research, some samples were found in a pool at the twilight zone of a cave (Fig. 1C) and in a pool in the dark zone of a cave, which was directly fed by the dripping water from stalactites (Fig. 1F). The favourite habitats of A. (C.) thailandica sp. nov. in the present study are likely to be phytotelmata or wet soils, as is already known from most species of Attheyella (subgenus Canthosella) (Reid 2001). At times, A. (C.) thailandica sp. nov. has been found together with Bryocyclop muscicola (Menzel, 1926); both are recognised as stygophile species and are widely distributed in Thailand (Watiroyram 2018; present study).

Discussion

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; P1P4 with two-segmented Enp (P1 Enp shorter than Exp, P4 Enp-1 much smaller than other legs of Enp-1); P2P4 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; P2P4 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 P2P4 legs. In contrast to SEA species, all American species, except A. (C.) acanthophora, show a higher number of setae on the P2P4 Enp-2 in both sexes. Females of these American species have three to four, three to five or two elements on P2P4, 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 P2P4, 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 1). (a) The P3 Enp-2 of a female in these species bears three elements, while it only bears two elements in A. (C.) lacustris, A. (C.) muscicola and A. (C.) silvicola and has only one seta in A. (C.) fluviatilis. (b). The P4 Enp in both sexes bears two apical setae, while it bears only one seta in all other individuals, except the male of A. (C.) silvicola, which has two setae. (c) The P2 Exp-1 in both sexes of these species has an enlarged outer spine, but this spine is normal in the other species; this characteristic is presumed to be a synapomorphy, which can be used to define closely-related species.

Table 1.

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
P2P4
– 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. 6B), but one-segmented in A. (C.) vietnamica. (b) The female caudal ramus of the new species has no spinules on the inner margin, but these spinules are present in A. (C.) vietnamica. (c) The new species has a dorsal seta located at three-quarters of the length of the caudal ramus, while this seta is located almost at the distal end in A. (C.) vietnamica. d) The male of the new species has two setae on the P3 Enp-3, while the male of A. (C.) vietnamica has one seta. e) A. (C.) thailandica sp. nov. exhibits a reduced Enp (especially P3) in both sexes, which are shorter in A. (C.) vietnamica. f).The morphology of P5 differs in several ways.

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.

A key to worldwide species of the subgenus Canthosella Chappuis, 1931

Females

(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

Males

(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

Acknowledgements

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.

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