taxonID	type	description	language	source
03F05637662AFFB0FF3A3921C8F24F36.taxon	description	(Figs. 3 – 7; Tables 3 and 4) urn: lsid: zoobank. org: act: AAF 98354 - B 04 D- 46 B 2 - B 217 - AFC 1 D 8 D 6 D 563 3.2.1. Synonym list Echinoderes sp. 1: Yamasaki et al. (2013); Dal Zotto et al. (2013).	en	Yamasaki, Hiroshi, Yoshida, Masa-aki, Jimi, Naoto, Hookabe, Natsumi, Sako, Misato, Kohtsuka, Hisanori, Fujimoto, Shinta (2024): Kinorhynch fauna from Oki Islands, with the description of a new Echinoderes species and its phylogenetic relationships within the family Echinoderidae. Zoologischer Anzeiger 313: 87-101, DOI: 10.1016/j.jcz.2024.08.006, URL: https://doi.org/10.1016/j.jcz.2024.08.006
03F05637662AFFB0FF3A3921C8F24F36.taxon	diagnosis	3.2.2. Diagnosis Echinoderes with middorsal acicular spines on segments 4 – 8; ventrolateral tubes on segment 2; lateroventral tubes on segment 5; lateroventral acicular spines on segments 6 – 9; midlateral tubes on segment 10; one type- 1 glandular cell outlets present middorsally each on segments 1 – 3, 5, and 7, two present middorsally on segments 10 and 11, one pairs present paradorsally on segments 4, 6, 8, and 9, sublaterally on segment 1, and ventromedially on segments 2 – 10; type- 2 glandular cell outlet absent; tergal extension narrowing abruptly into long acicular tips, constituting 5 – 7 % of trunk length; males with three pairs of penile spines of which middle pair very short; females with papillae in ventrolateral position on segment 7 and ventromedial position on segment 8, as well as with lateral terminal accessory spines.	en	Yamasaki, Hiroshi, Yoshida, Masa-aki, Jimi, Naoto, Hookabe, Natsumi, Sako, Misato, Kohtsuka, Hisanori, Fujimoto, Shinta (2024): Kinorhynch fauna from Oki Islands, with the description of a new Echinoderes species and its phylogenetic relationships within the family Echinoderidae. Zoologischer Anzeiger 313: 87-101, DOI: 10.1016/j.jcz.2024.08.006, URL: https://doi.org/10.1016/j.jcz.2024.08.006
03F05637662AFFB0FF3A3921C8F24F36.taxon	description	3.2.3. Etymology The specific name “ okiensis ” is derives from its type locality (Oki Islands).	en	Yamasaki, Hiroshi, Yoshida, Masa-aki, Jimi, Naoto, Hookabe, Natsumi, Sako, Misato, Kohtsuka, Hisanori, Fujimoto, Shinta (2024): Kinorhynch fauna from Oki Islands, with the description of a new Echinoderes species and its phylogenetic relationships within the family Echinoderidae. Zoologischer Anzeiger 313: 87-101, DOI: 10.1016/j.jcz.2024.08.006, URL: https://doi.org/10.1016/j.jcz.2024.08.006
03F05637662AFFB0FF3A3921C8F24F36.taxon	materials_examined	3.2.4. Material examined Holotype: Adult female (catalogue no. NSMT-Ki 52), collected on 21 June 2022 from rocky sediment sample at Onbe, Oki Islands, Japan (station Onbe- 1: 36 ◦ 9.185 ′ N, 133 ◦ 10.962 ′ E, 67 – 70 m depth), mounted in Fluoromount G ® on a H-S slide. Paratype: three adult males and one adult female (catalogue no. NSMT-Ki 53 – 56), and exoskeleton of one adult female used for DNA extraction (catalogue no. NSMT-Ki 57), all collected with the holotype; exoskeletons of two adult females used for DNA extraction (catalogue no. NSMT-Ki 58 – 59) collected on 19 August 2022 from rocky sediment sample at Onbe, Oki Islands, Japan (station Onbe- 2: 36 ◦ 9.218 ′ N, 133 ◦ 10.781 ′ E, 70 m depth); two adult males and one adult female (catalogue no. NSMT-Ki 60 – 62), collected on 20 May 2011 from muddy sand sample off Iejima Island, Okinawa, Japan (station 9 - 1: 26 ◦ 40.283 ′ N, 127 ◦ 45.433 ′ E, 138 m depth); three adult males and three adult females (catalogue no. NSMT-Ki 63 – 68), and exoskeleton of one adult female used for DNA extraction (catalogue no. ICHUM- 04291: the acronym of the museum collection was changed from ZIHU to ICHUM), collected on 20 May 2011 from muddy sand sample off Iejima Island, Okinawa, Japan (station 9 - 2: 26 ◦ 40.517 ′ N, 127 ◦ 45.333 ′ E, 124 m depth); all individually mounted in Fluoromount G ® on H-S slides. Non-type: one adult male and one adult female, collected with the holotype, mounted on aluminum stubs for SEM. 3.2.5. Type locality Onbe, Oki Islands, Japan (36 ◦ 9.185 ′ N, 133 ◦ 10.962 ′ E), 67 – 70 m depth.	en	Yamasaki, Hiroshi, Yoshida, Masa-aki, Jimi, Naoto, Hookabe, Natsumi, Sako, Misato, Kohtsuka, Hisanori, Fujimoto, Shinta (2024): Kinorhynch fauna from Oki Islands, with the description of a new Echinoderes species and its phylogenetic relationships within the family Echinoderidae. Zoologischer Anzeiger 313: 87-101, DOI: 10.1016/j.jcz.2024.08.006, URL: https://doi.org/10.1016/j.jcz.2024.08.006
03F05637662AFFB0FF3A3921C8F24F36.taxon	description	3.2.6. Description Adult with head, neck, and eleven trunk segments (Fig. 3 A, B, 4 A – D, 5 A). See Table 3 for the positions of cuticular structures. Table 4 indicates measurements. Head consists of retractable mouth cone and introvert (Fig. 5 A and B). Mouth cone comprised of inner and outer oral styles, however, detail of inner oral styles not observable in available specimens. Outer oral styles alternating in length: slightly longer in odd sector than in even sector. Outer oral style in sector 6 missing. Introvert with at least six rings of spinoscalids and one ring of trichoscalids (Fig. 5 B). Ring 01 with ten primary spinoscalids composed of one basal sheath and one distal end piece. Each basal sheath with proximal fringe composed of several projections between two long and slightly thicker lateral ones. Distal end pieces with proximal fringe with projections and long flexible spinose process, representing longest within all spinoscalids. Rings 02 and 04 with 10 spinoscalids; rings 03 and 05 with 20 spinoscalids. Each spinoscalid of rings 02 – 05 with basal sheath and end piece. Basal sheath of spinoscalids on ring 03 appeared to have a thick spinose process, whereas others composed of multiple thin and short spinose processes. Spinoscalids ring 06 not examined in detail, but with at least four spinoscalids distributed in section 4, 6, and 8. Six trichoscalids covered with long hairs on entire surface and each attached to trichoscalid plate. Number and arrangement of outer oral styles, and spinoscalids summarized in Fig. 6. Neck with 16 placids. Midventral placid wider than others, and remaining placids similar in width. Trunk with eleven segments; segment 1 consists of complete cuticular ring; segment 2 also consists of single cuticular ring, but with midventral scar, segments 3 – 11 consist of one tergal and two sternal plates (Fig. 4 E-N, 5 C-E, and 6). Thickened cuticle forms pachycyclus at anterior margin and beside midventral and tergosternal articulations of segments 2 – 10 (e. g., Fig. 4 A – H, J, K). Segment 1 with type- 1 glandular cell outlet in middorsal position (Fig. 3 A and 4 E). Additional pair of type- 1 glandular cell outlets in sublateral position (Fig. 3 B). Sensory spots present subdorsally and laterodorsally (Fig. 3 A, 4 E and 5 C). Non-bracteate cuticular hairs arising from perforation sites, covering whole segment (Fig. 5 C and D). Posterior edge with primary pectinate fringe consisted of very short fringe tips. Segment 2 with ventrolateral tubes (Fig. 3 B, 4 F and 5 D). Type- 1 glandular cell outlets, one in middorsal and one pair in ventromedial positions (Fig. 3 A, B, 4 E, F). One sensory spot present middorsally, two pairs laterodorsally, and one pair ventromedially (Fig. 3 A, B, 4 E, F, 5 C, D). Bracteate cuticular hairs from perforation sites covering whole segment except for muscular attachment sites in laterodorsal position as well as ventromedial – midventral area (Fig. 3 A, B, 5 C, D). Primary pectinate fringe similar to that on segment 1, but with slightly longer fringe tips especially on midlateral-lateroventral area. Segment 3 with one middorsal and one pair of ventromedial type- 1 glandular cell outlets (Fig. 3 A, B, 4 E, F). Sensory spots present in subdorsal and midlateral positions (Fig. 3 A, B, 4 E, F, 5 C). Cuticular hairs and primary pectinate fringe as on preceding segment. Segment 9 with lateroventral acicular spines (Fig. 3 A, B, 4 I, 7 A, D, E). One pair of type- 1 glandular cell outlets present each in paradorsal and ventromedial position (Fig. 3 A, B, 4 H, I). Sensory spots present in paradorsal, subdorsal, midlateral, and ventrolateral positions (Fig. 3 A, B, 4 I, 7 C). Small rounded sieve plate present in lateral accessory position (Fig. 3 B and 4 I). Cuticular hairs and primary pectinate fringe as on segment 5. Segment 10 with relatively long midlateral tubes (Figs. 3 and 4 L, N, 7 C – E). Two type- 1 glandular cell outlets present in middorsal position, and one pair in ventromedial position (Fig. 3). Sensory spots present subdorsally and ventrolaterally (Figs. 3 and 7 C). Cuticular hairs and primary pectinate fringe as on segment 5. Segment 11 with lateral terminal spines (Figs. 3 and 4 L – N, 5 A, 7 C – E). Males with three pairs of penile spines: most dorsal and most ventral penile spines long whereas middle penile spines short and visible only in SEM (Fig. 3 C, D, 4 N, 7 C, E). Females with lateral terminal accessory spines (Fig. 3 A, B, 4 L, 7 D). Two type- 1 glandular cell outlets present middorsally, and sensory spots present subdorsally (Figs. 3 and 7 E). Segment surface smooth and hairless. Posterior edge of tergal plate forming tergal extension narrowing abruptly into long acicular tips (Figs. 3 and 4 L – N, 5 A, 7 C, D). Segment 4 with middorsal acicular spine (Fig. 3 A, 4 E, G, and 5 A, C). One pair of type- 1 glandular cell outlets present each in paradorsal and ventromedial positions (Fig. 3 A, B, 4 F, H). Sensory spots present subdorsally and laterodorsally (Fig. 3 A, 4 E, G, and 5 C). Cuticular hairs and primary pectinate fringe as on segment 2. Segment 5 with middorsal acicular spine and lateroventral tubes (Fig. 3 A, B, 4 E – G, I, 5 E, 7 A). One pair of type- 1 glandular cell outlets present each in middorsal and ventromedial positions (Fig. 3 A, B, 4 F, H, I). Sensory spots present subdorsally, midlaterally, and ventromedially (Fig. 3 A, B, 4 E – G, I, 7 A). Cuticular hairs as on segment 2. Primary pectinate fringe with slightly longer fringe tips than those on segments 2 – 4. Segment 6 with middorsal and lateroventral acicular spines (Fig. 3 A, B, 4 G, I, 5 A, E, 7 A, B). One pair of type- 1 glandular cell outlets present each in paradorsal and ventromedial positions (Fig. 3 A, B, 4 H, I). Sensory spots present paradorsal, subdorsal, and midlateral positions (Fig. 3 A, B, 4 G). Cuticular hairs as on segment 2. Primary pectinate fringe as on segment 5. Segment 7 with middorsal and lateroventral acicular spines (Fig. 3 A, B, 4 G, I – K, 5 A, E, 7 A, B). Type- 1 glandular cell outlets, sensory spots, cuticular hairs, and primary pectinate fringe as on segment 5 (Fig. 3 A, B, 4 G – K, 5 E, 7 A, B). Females with papillae with tubular-shaped intracuticular substructure in ventrolateral position (Fig. 3 A, B, 4 I – K, 7 B). Segment 8 similar to segment 6, except for presence of ventromedial papillae in females (Fig. 3 A, B, 4 G – K, 5 E, 7 A – C). 3.2.7. Variation All the major morphological features — such as spines, tubes, glandular cell outlets, female papillae, and tergal extension shape — were congruent in all the Echinoderes specimens from the Oki Islands (stations Onbe- 1 and Onbe- 2) and off Iejima Island (stations 9 - 1 and 9 - 2). As shown in Table 4, morphometric data also overlapped between the two populations, and no significant differences were observed. Regarding the DNA sequences, 1776 bp of 18 S was determined for individuals from both the Oki Islands and off Iejima Island, and these sequences are completely identical. For 28 S, 3300 bp was determined for the specimen from off Iejima Island, while 3296 bp was determined for the specimen from the Oki Islands, with two undetermined internal regions (one representing 5 – 10 bp, and the other 25 – 30 bp). The 28 S sequences were compared after alignment, and except for the missing data regions, there were 7 nucleotide substitutions and 4 nucleotide insertions / deletions between the sequences of the two individuals. 3.2.8. Taxonomic remarks As mentioned in the previous section, Echinoderes individuals from the Oki Islands (stations Onbe- 1 and Onbe- 2) and off Iejima Island (stations 9 - 1 and 9 - 2) could not be distinguished morphologically. Regarding the DNA sequences, there is little data available for intraspecific variation in 18 S and 28 S sequences in Kinorhyncha. For example, in 18 S of Echinoderes songae Sørensen et al., 2020, two nucleotide substitutions were found in a comparable region of 1783 bp between Japanese (LC 831703) and Korean (GQ 229038) individuals. For 28 S, there is currently no comparable data available for intraspecific variation within any kinorhynch species. However, when comparing the differences between closely related species, for example, Echinoderes ohtsukai and Echinoderes rex (the two species are suggested to be closely related in both this study and a previous study using 18 S and COI (Randsø et al., 2019 )), there are three substitutions within 1772 bp in 18 S (LC 096961 for E. ohtsukai and LC 081126 for E. rex) and 52 substitutions as well as 9 bases insertions / deletions within 2206 bp in 28 S (LC 096963 for E. ohtsukai and LC 081131 for E. rex). This interspecific variation is significant compared to the differences between the Echinoderes individuals from the Oki Islands and off Iejima Island. Furthermore, the results of the phylogenetic analysis based on 18 S and 28 S sequences in this study also supported that the individuals from the two localities are closely related, forming a clade together (see the next section). Therefore, despite the geographic distance between the Oki Islands and Iejima Island, it is plausible to treat these individuals as the same species, E. okiensis sp. nov. In the family Echinoderidae, the spine / tube pattern of especially on segments 1 – 9 is the most easily recognized and is often the primary taxonomic trait used for the identification and classification of species (Yamasaki et al., 2020 b). However, the pattern in E. okiensis sp. nov. — the presence of middorsal acicular spines on segments 4 – 8, lateroventral / ventrolateral tubes on segment 2, lateroventral tubes / spines on segments 5 – 9 — is one of the most common spine / tube patterns in the family, shared with 31 congeners (Yamasaki et al., 2020 b). To distinguish species with such common spine / tube patterns, focusing on the pattern of type 1 / 2 glandular cell outlets and the shape of the tergal extension has recently gained attention as valuable taxonomic traits (Varney et al., 2019; Sørensen et al., 2020). These traits make it easy to distinguish E. okiensis sp. nov. from the other congeners.	en	Yamasaki, Hiroshi, Yoshida, Masa-aki, Jimi, Naoto, Hookabe, Natsumi, Sako, Misato, Kohtsuka, Hisanori, Fujimoto, Shinta (2024): Kinorhynch fauna from Oki Islands, with the description of a new Echinoderes species and its phylogenetic relationships within the family Echinoderidae. Zoologischer Anzeiger 313: 87-101, DOI: 10.1016/j.jcz.2024.08.006, URL: https://doi.org/10.1016/j.jcz.2024.08.006
