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Research Article
Taxonomy of the genus Elasmopus (Crustacea, Amphipoda) in Japan and South Korea, with description of a new species
expand article infoHiro Yoshimura, Chi-Woo Lee§, Ko Tomikawa
‡ Hiroshima University, Hiroshima, Japan
§ Nakdonggang National Institute of Biological Resources, Sangju, Republic of Korea
Open Access

Abstract

To advance the limited understanding of the East Asian Elasmopus fauna, field surveys were conducted in Japan and South Korea. In the present study, a new species, E. lumbiniger sp. nov., is described. The new species is distinguished from its congeners by a strongly projected posteroventral corner of the epimeral plate 3, long slender setae on the anterodistal corner of the male gnathopod 2 carpus, a mid-palmar ridge on the male gnathopod 2 propodus, and a smooth posterior margin on the basis of pereopods 5–7. In addition, E. koreanus is recorded in Japan for the first time, and the known distribution range of E. mukuinu has been significantly extended. The nucleotide sequences of the mitochondrial cytochrome c oxidase subunit I (COI) of these species were determined, and the genetic distances among Elasmopus species were provided. A key to the species of Elasmopus found in East Asia is also provided.

Key words:

COI, East Asia, genetic distance, Kimura two-parameter distance, morphology, p-distance

Introduction

The genus Elasmopus Costa, 1853 is distributed in the shallow waters of tropical and temperate regions worldwide (Lowry and Hughes 2009; Vader and Krapp-Schickel 2012). To date, 124 Elasmopus species have been described (Vader and Krapp-Schickel 2012; Alves et al. 2016; Horton et al. 2025; Yoshimura and Tomikawa 2025). Of these, 13 species of Elasmopus are currently known from East Asia (Table 1).

Table 1.

Occurrence records of Elasmopus species in East Asia.

Species Occurrence in East Asia Notes References
E. alkhiranensis Myers & Momtazi, 2015 South China Sea, China Ren (2012) recorded this species as E. pectenicrus (Bate, 1862); however, it was referred to E. alkhiranensis by Myers and Momtazi (2015). Ren (2012); Myers and Momtazi (2015)
E. hawaiensis Schellenberg, 1938 South China Sea, China - Ren (2012)
E. hooheno Barnard, 1970 South China Sea, China - Ren (2012)
E. japonicus Stephensen, 1932 Philippine Sea, Japan - Stephensen (1932)
E. koreanus Kim & Kim, 1991 Sea of Japan (East Sea), South Korea, and Japan; Pacific Ocean, Japan - Kim and Kim (1991); Kim et al. (2023); Shin et al. (2025); the present study
E. lumbiniger sp. nov. Sea of Japan, Japan (East Sea); Philippine Sea, Japan; Pacific Ocean, Japan - present study
E. mukuinu Sir & White, 2022 Philippine Sea, Japan - Sir and White (2022); present study
E. nanshaensis Ren, 1998 South China Sea, China - Ren (2012)
E. nkjaf Nakamura, Nakano, Ota & Tomikawa, 2019 Philippine Sea, Japan - Nakamura et al. (2019)
E. projectus Yoshimura & Tomikawa, 2025 Sea of Japan (East Sea), Japan; Pacific Ocean, Japan - Yoshimura and Tomikawa (2025)
E. rapax Costa, 1853 South China Sea, China; Sea of Japan (East Sea), South Korea - Shin et al. (2005); Ren (2012); Shin et al. (2025)
E. spinicarpus Berents, 1983 South China Sea, China - Ren (2012)
E. spinidactylus Chevreux, 1907 South China Sea, China - Ren (2012)
E. spinimanus Walker, 1904 South China Sea, China - Ren (2012)

In Japan, E. japonicus and E. projectus have been recorded from temperate coastal areas, and E. mukuinu and E. nkjaf from subtropical zones (Stephensen 1932; Nakamura et al. 2019; Sir and White 2022; Yoshimura and Tomikawa 2025). Two species, E. koreanus and E. rapax, have been recorded from South Korean waters (Kim and Kim 1991; Shin et al. 2005; Kim et al. 2023; Shin et al. 2025). However, many regions in Japan and South Korea remain taxonomically unstudied with regard to Elasmopus. The presence of several unrecorded or undescribed species of Elasmopus has been suggested (Hirayama 1995; Ariyama 2022).

To clarify the Elasmopus fauna in Japan and South Korea, we conducted field surveys at multiple sites. We found E. koreanus and E. mukuinu in temperate regions of Japan, where they had not been previously recorded, as well as an undescribed species, E. lumbiniger sp. nov. Additionally, we sequenced the mitochondrial cytochrome c oxidase subunit I (COI) gene of the collected samples and calculated the genetic distances among the Elasmopus species.

Materials and methods

Field sampling

The specimens were collected from the intertidal and shallow subtidal zones during low tide, depth range 0–1 m. The sampling sites are shown in Fig. 1. While still alive, some specimens were photographed using a digital camera (EOS M6; Canon, Tokyo, Japan) with a lens (LAOWA 65 mm F2.8 Macro Photo Lens; Venus Optics, Heifei, China). After photographs were taken, some specimens were fixed and preserved in 99% ethanol while others were frozen, fixed, and preserved in propylene glycol.

Figure 1. 

Map showing collecting sites of the examined specimens. Slashes within boxes indicate that two species were collected from the same site. A. Kannonzaki Coast, Kanagawa, the type locality of Elasmopus lumbiniger sp. nov.; B. Arasaki Coast, Kanagawa; C. Uchiura Beach, Chiba; D. Shirahama, Chiba; E. Ono Beach, Shizuoka; F. Ôura Beach, Shikinejima Island, Tokyo; G. Yakengahama Beach, Hachijojima Island, Tokyo; H. Uradome Beach, Tottori; I. Takurazaki, Wakayama; J. Kaichu Doro, Okinawa Island, Okinawa, the type locality of E. mukuinu; K. Songjeonggudeokpo-gil, Busan.

Morphological examinations

All appendages were dissected in 50% ethanol and mounted in gum-chloral medium on glass slides under a stereomicroscope (SZ61; Olympus, Tokyo, Japan). The specimens were examined using a light microscope (Eclipse Ni-U; Nikon, Tokyo, Japan) and illustrated with the aid of a camera lucida (Y-IDT Drawing Tube; Nikon, Tokyo, Japan). Body length, measured from the rostrum tip to the base of the telson along the dorsal curvature, was recorded to the nearest 0.1 mm. Some slide preparations were photographed using a device that integrated a digital camera (EOS M6) with an imaging lens (EF-M 55-200 mm F4.5–6.3 IS STM; Canon, Tokyo, Japan) and an objective lens (UPlan Fl 10x/0.30; Olympus, Tokyo, Japan). The examined specimens are deposited in the National Museum of Nature and Science, Tsukuba (NSMT), and Nakdonggang National Institute of Biological Resources (NNIBR).

DNA sequencing

Genomic DNA was extracted from the pereonites and pleonites muscles of the specimens, using the protocol described by Tomikawa et al. (2014). The primer set used for polymerase chain reaction (PCR) and cycles sequencing (CS) was LCO1490 and HCO2198 (Folmer et al. 1994). PCR and CS mixtures were prepared following Tomikawa et al. (2014). PCR was performed as follows: enzyme activation for 5 min at 94 °C; 35 cycles of 30 s at 94 °C, 60 s at 50 °C, and 90 s at 72 °C; and final extension for 6 min at 72 °C. For samples with poor gene amplification efficiency under the above condition, the annealing temperature was adjusted from 50 to 42 °C. CS was performed as follows: 1 min at 96 °C; 25 cycles of 10 s at 96 °C, 5 s at 50 °C, and 4 min at 60 °C. The obtained sequences were assembled and edited using MEGA 11 (Tamura et al. 2021). The DNA sequence has been deposited with the International Nucleotide Sequence Database (INSD) through the DNA Data Bank of Japan (DDBJ). The INSD accession numbers are LC872568LC872582 (Table 2).

Table 2.

Specimen details and GenBank accession numbers used in the present study. Newly obtained sequences are indicated by an asterisk.

Species Sample number Locality Coordinates (decimal degrees) GenBank accession number
E. koreanus NSMT-Cr 33060 Shirahama, Chiba, Japan 34.9031°N, 139.8862°E *LC872568
NSMT-Cr 33062 Uradome, Tottori, Japan 35.5915°N, 134.3220°E *LC872569
NNIBRIV137288 Songjeonggudeokpo-gil, Busan, South Korea 35.1682°N, 129.1975°E *LC872570
E. lumbiniger sp. nov. NSMT-Cr 33064 Kannonzaki Coast, Kanagawa, Japan 35.2592°N, 139.7436°E *LC872571
NSMT-Cr 33072 Arasaki Coast, Kanagawa, Japan 35.1947°N, 139.5996°E *LC872572
NSMT-Cr 33073 Uchiura Beach, Chiba, Japan 35.1262°N, 140.1893°E *LC872573
NSMT-Cr 33074 Shirahama, Chiba, Japan 34.9031°N, 139.8862°E *LC872574
NSMT-Cr 33076 Takurazaki, Wakayama, Japan 34.2671°N, 135.0605°E *LC872575
NSMT-Cr 33078 Uradome, Tottori, Japan 35.5921°N, 134.3198°E *LC872576
E. mukuinu NSMT-Cr 33079 Okinawa Island, Okinawa, Japan 26.3324°N, 127.9222°E *LC872577
NSMT-Cr 33080 Ono Beach, Shizuoka, Japan 34.6329°N, 138.8975°E *LC872578
NSMT-Cr 33082 Shikinejima Island, Tokyo, Japan 34.3307°N, 139.2083°E *LC872579
NSMT-Cr 33084 Hachijôjima Island, Tokyo, Japan 33.1007°N, 139.7696°E *LC872580
E. nkjaf KUZ Z1864 Miyako Island, Okinawa, Japan 24.7997°N, 125.3342°E LC215812
KUZ Z1862 Miyako Island, Okinawa, Japan 24.7997°N, 125.3342°E LC215813
E. projectus NSMT-Cr 32973 Kannonzaki Coast, Kanagawa, Japan 35.2592°N, 139.7436°E LC851065
NSMT-Cr 32974 Kannonzaki Coast, Kanagawa, Japan 35.2592°N, 139.7436°E *LC872581
NSMT-Cr 32980 Uradome, Tottori, Japan 35.5921°N, 134.3198°E *LC872582
E. rapax SFAM20-013 Somewhere along the west coast of Portugal - KF369125
SFC18-003 Viana do Castelo, Minho, Portugal 41.70°N, 8.85°W KX224027
SFAM12-002 Viana do Castelo, Minho, Portugal 41.70°N, 8.85°W KX224028
SFAM20-001 Viana do Castelo, Minho, Portugal 41.70°N, 8.85°W KX224029
SFAM20-002 Viana do Castelo, Minho, Portugal 41.70°N, 8.85°W KX224030
SFAM20-003 Viana do Castelo, Minho, Portugal 41.70°N, 8.85°W KX224031

Genetic distances

Genetic distances within and among E. koreanus, E. lumbiniger sp. nov., E. mukuinu, E. nkjaf, E. projectus, and E. rapax were calculated. Nucleotide sequences of E. nkjaf, E. projectus, and E. rapax were obtained from NCBI database (https://www.ncbi.nlm.nih.gov/) (Table 2). Sequence alignment was performed using the MUSCLE algorithm implemented in MEGA 11. Genetic divergences were estimated based on uncorrected pairwise distance (p-distance) and Kimura’s two-parameter model (K2P) (Kimura 1980).

Results

Systematics

Family Maeridae Krapp-Schickel, 2008

Genus Elasmopus Costa, 1853

Elasmopus koreanus Kim & Kim, 1991

Figs 2, 3, 4, 5 [New Japanese name: Kourai-iso-yokoebi]

Elasmopus koreanus Kim & Kim, 1991: 329, figs 6, 7.

Material examined.

• Male, 10.1 mm (NSMT-Cr 33059), Kannonzaki Coast, Yokosuka, Kanagawa, Japan (35.2590°N, 139.7429°E) (Fig. 1A), shallow subtidal zone, rocky shore, among holdfast of the brown alga Sargassum fusiforme, 1 March 2022, H. Yoshimura coll.; • male, 8.3 mm (NSMT-Cr 33060), Shirahama, Minamiboso, Chiba, Japan (34.9031°N, 139.8862°E) (Fig. 1D), intertidal zone, rocky shore, 5 June 2023, H. Ogawa coll.; • male, 8.3 mm (NSMT-Cr 33061), Oura Beach, Shikinejima, Tokyo, Japan (34.3307°N, 139.2083°E) (Fig. 1F), shallow subtidal zone, rocky shore, among the red algae, 3 August 2024, H. Yoshimura coll.; • male, 6.4 mm (NSMT-Cr 33062: Fig. 2A), ovigerous female, 6.2 mm (NSMT-Cr 33063: Fig. 2B), Uradome Beach, Iwami, Tottori, Japan (35.5915°N, 134.3220°E) (Fig. 1H), shallow subtidal zone, rocky shore, among holdfast of the red alga Hypnea sp., 16 June 2024, H. Ogawa, Y. Mukaida, M. Ooga, and H. Yoshimura coll.; • male, 5.0 mm (NNIBRIV137288), Songjeonggudeokpo-gil, Haeundae-gu, Busan, South Korea (35.1682°N, 129.1975°E) (Fig. 1K), tide pool, rocky shore, among the brown alga Sargassum thunbergii, 26 September 2024, H. Yoshimura, C.W. Lee and K. Tomikawa coll.

Figure 2. 

Elasmopus koreanus. A. Male, 6.4 mm (NSMT-Cr 33062), lateral view; B. Female, 6.2 mm (NSMT-Cr 33063), lateral view. Scale bar: 1.0 mm.

Type locality.

Sadong, Ulleungdo Island, South Korea (37.472°N, 130.888°E; assumed from Kim and Kim 1991: fig. 1).

Diagnosis.

Epimeral plate 3 posteroventral corner without distinct projection. Antenna 1 peduncles bearing slender setae on lateral and medial margins, with bi-articulate accessory flagellum. Antenna 2 flagellum articles distinctly broader than long. Mandibular palp article 3 short. Male gnathopod 2 propodus posterior margin with a few clusters of slender setae, palmar margin with subtriangular process near hinge of dactylus, mid-palmar margin with subrectangular hump. Length of setae on coxae 2 and 3 shorter than width of each coxa. Pereopods 5–7 basis posterior margin smooth, without long setae. Telson cleft, border than long; lobe trapezoid, with shallow incision at apex, each lobe bearing one plumose and two or three robust setae apically.

Description of male

(NSMT-Cr 33062). Body (Fig. 2A): smooth, not carinate, with a few short setae dorsally on pereon and abdomen. Head: eyes oval; lateral cephalic lobe broad, anteroventral margin with notch; dorsal surface with a few short setae. Epimeral plate 3 (Fig. 3A): ventral margin with seven robust setae, posterior margin with five short setae, posteroventral corner without distinct projection.

Figure 3. 

Elasmopus koreanus, male, 6.4 mm (NSMT-Cr 33062). A. Left epimeral plate 3, lateral view; B. Right accessory flagellum, medial view; C. Left antenna 2, dorsal view; D. Incisor, lacinia mobilis, accessory setal row and molar process of left mandible, medial view; E. Right mandible, medial view; F. Telson, dorsal view. Scale bars: 0.3 mm (A, C); 0.1 mm (B, D–F).

Antenna 1 : length 0.6 × body length; peduncular articles 1–3 with slender setae on lateral and medial margins; accessory flagellum (Fig. 3B) bi-articulate, reaching 2/3 of primary flagellar article 1, accessory flagellar article 2 tiny; primary flagellum 21-articulate, with slender setae. Antenna 2 (Fig. 3C): length 0.4 × antenna 1; peduncular articles 4 and 5 almost same length, with slender setae on lateral and medial margins; flagellum 8-articulate, articles 1–5 shorter than wide, with slender setae.

Mandible (Fig. 3D, E) with left and right incisors with two and four teeth, respectively; both left and right lacinia mobilis with five teeth; accessory setal row consisting of four setae on each of left and right mandibles; molar process well developed, triturative; palp well developed, tri-articulate; palp article 1 without setae, article 2 with six setae, article 3 falcate, length 2.5 × width.

Gnathopod 1 (Fig. 4A): subchelate; coxa anteroventral corner weakly produced, ventral margin with long and short setae; basis with long setae on posterior margin and medial surface; carpus subequal in length to propodus, with slender seta on anterodistal corner, with dense setae on posterior margin and medial surface; propodus with three and six clusters of setae on anterior margin and medial surface, respectively, posterior margin with row of slender and robust setae, palmar margin almost transverse, minutely serrate, with rows of robust setae on medial and lateral palmar submargins. Gnathopod 2 (Figs 4B, 5B): subchelate; coxa with long and short setae on ventral margin, length of the longest seta 0.6 × width of coxa; basis with long setae on posterior margin; carpus with dense setae on posterior margin; propodus with four setae on anterolateral submargin, posterior margin with nine clusters of slender setae, palmar margin with subrectangular hump at midpoint and robust seta at proximal end where tip of dactylus contacts, lateral palmar margin with subtriangular hump near hinge of dactylus.

Figure 4. 

Elasmopus koreanus, A, B. Male, 6.4 mm (NSMT-Cr 33062); C, D. Female, 6.2 mm (NSMT-Cr 33063). A. Right gnathopod 1, medial view; B. Left gnathopod 2, lateral view; C. Right gnathopod 1, medial view; D. Right gnathopod 2, medial view. Scale bars: 0.3 mm.

Figure 5. 

Elasmopus koreanus, ontogenetic morphological change in male gnathopod 2, merus to dactylus of right gnathopod 2, medial view. A. Male, 10.1 mm (NSMT-Cr 33059); B. Male, 6.4 mm (NSMT-Cr 33062); C. Male, 5.0 mm (NNIBRIV137288). Scale bars: 0.3 mm.

Telson (Fig. 3F): broader than long, cleft for 60% of length; lobe trapezoid, with shallow incision at apex, each lobe bearing one plumose and three robust setae apically, lateral margin of each lobe with none or one plumose setae.

Description of female

(NSMT-Cr 33063), sexually dimorphic characters. Gnathopod 1 (Fig. 4C): propodus anterior margin with one single and three clusters of setae, medial surface with five clusters of setae. Gnathopod 2 (Fig. 4D): propodus with nine and eight clusters of setae on anteromedial margin and medial surface, respectively, palmar margin almost straight, with minutely serrate on the distal one third of the palmar margin. Oostegites on gnathopod 2 and pereopods 3–5, slender with long setae.

Variations.

Mandible : left and right incisors with two or three and three or four teeth, respectively; lacinia mobilis with four or five teeth. Gnathopod 2: length of setae on coxa 0.4–0.9 × width of coxa. Pereopod 3: length of setae on coxa 0.6–0.9 × width of coxa. Uropod 3: the length-to-width ratio of inner ramus 1.5–2.5. Telson: each lobe bearing two or three robust setae apically.

As body length increases, male gnathopod 2 shows morphological changes (Fig. 5A–C): in adult males (features of immature males in parentheses), propodus elongate (semioval), hump on palmar margin strongly produced (weakly produced), mid-palmar margin with long ridge (without ridge), posterior margin without robust seta (with a robust seta at distal end of posterior margin).

Coloration in life.

Eyes black; antennae 1 and 2 peduncular articles brown, flagellum articles have two black patterns on a white background; body laterally brown, dorsally black, with small white dot patterns on head, pereon, and abdomen. Males and females have similar body coloration.

Distribution.

South Korea: Busan (present study); Ulleungdo Island in Gyeongsangbuk-do (Kim and Kim 1991; Shin et al. 2025); Chujado Islands in Jeju-do (Kim et al. 2023). Japan: Chiba, Kanagawa, and Tottori; Shikinejima Island in Tokyo (present study) (Fig. 1).

Remarks.

Our specimens obtained from South Korea and Japan are well identical to the original description of Elasmopus koreanus by Kim and Kim (1991). A comparison of E. koreanus from Japan and South Korea showed no remarkable morphological differences other than their ontogenetic changes. The intraspecific genetic distances of COI between Japanese and Korean specimens of this species were at most 0.024 (p-distance) and 0.025 (K2P), respectively (Tables 3, 4). These values were not particularly large compared to the intraspecific genetic distances of the other Elasmopus species examined in this study (Tables 3, 4). Therefore, we concluded that E. koreanus from Japan and South Korea should be regarded as conspecific, since they show no significant morphological or genetic differences that would support their separation as distinct species.

Table 3.

Uncorrected pairwise distances (p-distances) among Elasmopus species showing minimum–maximum values (mean value). “n” indicates the number of specimens, and “loc.” Indicates the number of collecting sites.

E. koreanus (n = 3, loc. = 3) E. lumbiniger sp. nov. (n = 6, loc. = 6) E. mukuinu (n = 4, loc. = 4) E. nkjaf (n = 2, loc. = 1) E. projectus (n = 3, loc. = 2) E. rapax (n = 6, loc. = 2)
E. koreanus 0.002–0.024 (0.016)
E. lumbiniger sp. nov. 0.198–0.205 (0.201) 0.002–0.043 (0.017)
E. mukuinu 0.202–0.207 (0.204) 0.167–0.184 (0.173) 0.003–0.050 (0.027)
E. nkjaf 0.190–0.193 (0.192) 0.185–0.191 (0.188) 0.178–0.182 (0.180) 0.006 (-)
E. projectus 0.178–0.182 (0.181) 0.158–0.170 (0.163) 0.164–0.176 (0.172) 0.187–0.191 (0.189) 0.008–0.024 (0.017)
E. rapax 0.188–0.191 (0.190) 0.187–0.198 (0.190) 0.173–0.187 (0.178) 0.160–0.167 (0.162) 0.182–0.193 (0.186) 0.000–0.003 (0.001)
Table 4.

Kimura two-parameter distances (K2P distances) among Elasmopus species showing minimum–maximum values (mean value). “n” indicates the number of specimens, and “loc.” Indicates the number of collecting sites.

E. koreanus (n = 3, loc. = 3) E. lumbiniger sp. nov. (n = 6, loc. = 6) E. mukuinu (n = 4, loc. = 4) E. nkjaf (n = 2, loc. = 1) E. projectus (n = 3, loc. = 2) E. rapax (n = 6, loc. = 2)
E. koreanus 0.002–0.025 (0.017)
E. lumbiniger sp. nov. 0.233–0.244 (0.238) 0.002–0.044 (0.017)
E. mukuinu 0.239–0.247 (0.243) 0.192–0.216 (0.201) 0.003–0.052 (0.028)
E. nkjaf 0.222–0.226 (0.224) 0.215–0.222 (0.219) 0.205–0.211 (0.208) 0.006 (-)
E. projectus 0.207–0.213 (0.210) 0.181–0.198 (0.189) 0.189–0.207 (0.200) 0.218–0.224 (0.221) 0.008–0.025 (0.018)
E. rapax 0.217–0.224 (0.221) 0.219–0.235 (0.224) 0.200–0.219 (0.206) 0.211–0.226 (0.217) 0.186–0.196 (0.189) 0.000–0.003 (0.001)

Elasmopus koreanus is most similar to E. mutatus Barnard, 1962, with the following five features: 1) epimeral plate 3 posteroventral corner without distinct projection; 2) male gnathopod 2 propodus mid-palmar margin with subrectangular hump; 3) male gnathopod 2 propodus medial surface without dense setae; 4) pereopods 5–7 basis smooth, without long setae; and 5) telson broader than long. Kim and Kim (1991) distinguished E. koreanus from E. mutatus based on four morphological characters: 1) antennae of E. koreanus bear fewer setae than those of E. mutatus; 2) male gnathopod 2 palmar margin ridge where tip of dactylus contacts lacks a robust seta in E. koreanus, whereas it bears a robust seta in E. mutatus; 3) posterior margin of male gnathopod 2 propodus covered with fewer setae in E. koreanus than in E. mutatus; and 4) uropod 3 inner ramus broader in E. koreanus than in E. mutatus. However, all examined male specimens in the present study possessed a robust seta on the palm of the gnathopod 2 at the point where the tip of the dactylus contacts. This suggests that either Kim and Kim (1991) overlooked this robust seta, or that the presence or absence of this robust seta represents intraspecific variation in E. koreanus. Therefore, the presence or absence of this robust seta is not a diagnostic character distinguishing E. koreanus from E. mutatus. Additionally, regarding the broadness of the uropod 3 inner ramus, the length-to-width ratio in E. koreanus ranged from 1.5 to 2.5. In E. mutatus, this ratio was 2.1, which fell within the intraspecific variation range of E. koreanus. Thus, E. koreanus and E. mutatus cannot be distinguished based on the broadness of the uropod 3 inner ramus. On the other hand, we propose newly found taxonomic features to distinguish the two species: 1) antennae of E. koreanus bear fewer setae than those of E. mutatus; 2) posterior margin of male gnathopod 2 propodus covered with fewer setae in E. koreanus than in E. mutatus; and 3) the longest seta on coxae 2 and 3 shorter than the width of each coxa in E. koreanus, whereas longer in E. mutatus.

Elasmopus lumbiniger sp. nov.

Figs 6, 7, 8, 9, 10, 11, 12 [New Japanese name: Sewata-iso-yokoebi]

Type material.

Holotype : • male, 9.8 mm (NSMT-Cr 33064), Kannonzaki Coast, Yokosuka, Kanagawa, Japan (35.2592°N, 139.7436°E) (Fig. 1A), shallow subtidal zone, rocky shore, among holdfast of the brown alga Eisenia bicyclis, 4 February 2023, H. Yoshimura coll. Paratypes: • male, 10.4, 9.9, 8.5 mm (NSMT-Cr 33065–33067), ovigerous female, 8.7 mm (NSMT-Cr 33068), same data as for the holotype; • male, 9.6, 7.1 mm (NSMT-Cr 33069, 33070), female, 5.9 mm (NSMT-Cr 33071), Kannonzaki Coast, Yokosuka, Kanagawa, Japan (35.2592°N, 139.7432°E) (Fig. 1A), shallow subtidal zone, rocky shore, among holdfast of the brown alga Sargassum sp., 22 April 2023, H. Yoshimura coll.; • male, 7.0 mm (NSMT-Cr 33072), Arasaki Coast, Yokosuka, Kanagawa, Japan (35.1947°N, 139.5996°E) (Fig. 1B), shallow subtidal zone, rocky shore, among the red alga Corallina sp., 2 September 2023, H. Yoshimura coll.; • male, 13.8 mm (NSMT-Cr 33073: Fig. 6A), Uchiura Beach, Kamogawa, Chiba, Japan (35.1262°N, 140.1893°E) (Fig. 1C), shallow subtidal zone, rocky shore, among the red alga Corallina sp., 18 June 2022, H. Yoshimura coll.; • male, 10.0 mm (NSMT-Cr 33074), Shirahama, Minamiboso, Chiba, Japan (34.9031°N, 139.8862°E) (Fig. 1D), intertidial zone, rocky shore, 5 June 2023, H. Ogawa coll.; • female, 12.6, 10.3 mm (NSMT-Cr 33075: Fig. 6B, NSMT-Cr 33076), Takurazaki, Wakayama, Wakayama, Japan (34.2671°N, 135.0605°E) (Fig. 1I), shallow subtidal zone, rocky shore, under the stone, 10 May 2024, H. Yoshimura coll.; • male, 10.0 mm (NSMT-Cr 33077), female, 7.9 mm (NSMT-Cr 33078), Uradome Beach, Iwami, Tottori, Japan (35.5921°N, 134.3198°E) (Fig. 1H), shallow subtidal zone, rocky shore, among sporophyll of the brown alga Undaria pinnatifida lying on the sea bottom, 16 June 2024, H. Ogawa, Y. Mukaida, M. Ooga, and H. Yoshimura coll.

Figure 6. 

Elasmopus lumbiniger sp. nov. A. Paratype male, 13.8 mm (NSMT-Cr 33073), lateral view; B. Paratype female, 12.6 mm (NSMT-Cr 33075), lateral view. Scale bar: 1.0 mm.

Diagnosis.

Epimeral plate 3 posteroventral corner produced into well-developed acute spine. Antenna 1 peduncles bearing slender setae on lateral and medial margins, with tri-articulate accessory flagellum. Mandibular palp article 3 short. Gnathopod 2 carpus anterodistal corner with slender setae, length of the longest seta reaching 0.8–1.1 and 0.9–1.1 × width of carpus in male and female. Male gnathopod 2 propodus posterior margin and inner surface with dense slender setae, length of the longest seta reaching 0.9 × width of propodus, palmar margin with trapezoidal hump near hinge of dactylus, mid-palmer margin with ridge. Pereopods 5–7 basis posterior margin smooth, without long setae. Telson cleft, border than long; lobe trapezoid, with shallow incision at apex, each lobe bearing one plumose and 4–6 robust setae apically.

Description of male

(holotype, NSMT-Cr 33064). Body (Fig. 7A): smooth, not carinate, with a few short setae dorsally on pereon and abdomen. Head: eyes oval; lateral cephalic lobe broad, anteroventral margin with notch; dorsal surface with a few short setae. Epimeral plates 1–3 (Fig. 7B–D): each ventral margin with one cluster and one single robust seta, two clusters and one single robust seta, and two clusters and five robust setae, respectively; each posterior margin with three or four, four and four short setae, respectively; posteroventral corner of epimeral plates 1 and 2 weakly produced, epimeral plate 3 produced into well-developed acute spine.

Figure 7. 

Elasmopus lumbiniger sp. nov. holotype male, 9.8 mm (NSMT-Cr 33064). A. Body, lateral view; B. Right epimeral plate 1, lateral view; C. Right epimeral plate 2, lateral view; D. Right epimeral plate 3, lateral view. Scale bars: 1.0 mm (A); 0.5 mm (B–D).

Antenna 1 (Fig. 8A): length 0.55 × body length; peduncular articles 1–3 in length ratio of 1.1: 1.0: 0.6, with slender setae on lateral and medial margin; peduncular article 1 with three robust setae on posteroproximal margin, robust seta on posterodistal corner; accessory flagellum tri-articulate, accessory flagellar article 3 tiny; primary flagellum 34-articulate, with slender setae. Antenna 2 (Fig. 8B): length 0.5 × antenna 1; peduncular article 2 with six short spines on anterodistal corner, gland cone of peduncular article 2 exceeding distal margin of peduncular article 3; peduncular article 4 slightly longer than 5, with slender setae on lateral and medial margins; flagellum 13-articulate, with slender setae.

Figure 8. 

Elasmopus lumbiniger sp. nov. A, B, D–J. Holotype male, 9.8 mm (NSMT-Cr 33064); C. Paratype male, 9.9 mm (NSMT-Cr 33066). A. Right antenna 1, medial view; B. Left antenna 2, lateral view; C. Upper lip, anterior view; D, E. Incisor, lacinia mobilis, accessory setal row and molar process of left and right mandible, medial view; F. Left mandibular palp, medial view; G. Lower lip, ventral view; H. Right maxilla 1, dorsal view; I. Right maxilla 2, dorsal view; J. Maxilliped, dorsal view. Scale bars: 0.5 mm (A, B); 0.1 mm (C–J).

Mouthparts. As holotype upper lip missing, that of paratype (male, 9.9 mm, NSMT-Cr 33066) illustrated. Upper lip (Fig. 8C) with rounded anterior margin, bearing fine setae. Mandible (Fig. 8D–F) with left and right incisors two and four teeth, respectively; left lacinia mobilis five teeth and right lacinia mobilis four teeth; accessory setal row three setae on the left and four setae on the right; molar process well developed, triturative; palp (Fig. 8F) well developed, tri-articulate; palp article 1 without setae, article 2 with ten setae, article 3 falcate, length 3.0 × width. Lower lip (Fig. 8G) with outer lobes laterally expanded, bearing apical setae; inner lobes ovate, apically covered with setae. Maxilla 1 (Fig. 8H) with inner plate bearing two plumose setae apically; outer plate with seven robust dentate setae; palp bi-articulate, article 1 marginally bare, article 2 with numerous apical and subapical setae, outer margin bare. Maxilla 2 (Fig. 8I) with inner plate bearing long apical setae, with setae on outer and inner margins; outer plate bearing long apical setae, with setae on outer margin. Maxilliped (Fig. 8J) with inner plate bearing plumose setae apically; outer plate obovate, with spatula plumose setae on apical and inner margins, reaching more than half of palp article 2; palp article 1 with two setae on inner margin, with seta on outer margin; palp article 2 slender, lined with setae on inner margin, outer margin bare; nail of palp article 4 distinct.

Gnathopod 1 (Fig. 9A–D): subchelate; coxa anteroventral corner weakly produced, anterior margin weakly concave, ventral margin with long and short setae; basis with long setae on posterior margin and medial surface; carpus subequal in length to propodus, with slender setae on anterodistal corner, with dense pinnate sided setulated setae on medial surface and posterior margin; propodus with six clusters of setae on anterior margin, with seven clusters of setae on medial surface; palmar margin (Fig. 9C, D) almost transverse, minutely serrate, with rows of robust setae on medial and lateral palmar submargins, palmar corner with two long robust setae; dactylus with slender seta on anteroproximal margin, with six small setae on lateral margin, with spatulate seta and slender seta laterally at the base of nail. Gnathopod 2 (Fig. 9E–G): subchelate; coxa with long and short setae on ventral margin, of which two long setae retrorse; basis with long setae on posterior margin; merus distoventral corner subquadrate bearing long setae; carpus with slender setae on anterodistal corner, length of these setae 0.9 × width of carpus, with dense setae on posterior margin; propodus with five clusters, two pairs and two single setae on anterolateral submargin, length of these setae 0.4 × width of propodus, anteromedial submargin with eight clusters of setae; lateral palmar margin weakly concave with trapezoidal hump (Fig. 9G) on near insertion of dactylus, with one clusters, three pairs and one single setae on lateral surface, with six and seven robust setae on lateral and medial margins, respectively; mid-palmar margin with ridge; posterior margin with row of plumose setae, the proximal seta longer than the distal seta, length of the longest seta 0.9 × width of propodus; dactylus strongly curved, subacute, length 0.6 × length of propodus, with seta on anteroproximal margin.

Figure 9. 

Elasmopus lumbiniger sp. nov. holotype male, 9.8 mm (NSMT-Cr 33064). A. Right gnathopod 1, medial view; B. Left gnathopod 1, lateral view; C. Palmar margin of right gnathopod 1 propodus, medial view; D. Palmar margin of left gnathopod 1 propodus, lateral view; E. Right gnathopod 2, medial view; F. Left gnathopod 2, lateral view, coxal gill omitted; G. Hump on right gnathopod 2 propodus, medial view. Gnathopod 2 coxal gill omitted. Scale bars: 0.5 mm (A, B, E, F); 0.1 mm (C, D, G).

Pereopod 3 (Fig. 10A): coxa longer than broad, ventral margin with numerous short and a few long setae, of which long seta retrorse; posterior margins of basis with long slender and robust setae, merus with slender setae, carpus and propodus with robust and slender setae; dactylus with plumose seta on anteroproximal margin, with one robust and two slender setae at the base of nail. Pereopod 4 (Fig. 10B): coxa expanded with posterior concavity, ventral margin arched bearing numerous short and a few long setae, of which long seta retrorse; posterior margins of basis with long slender and robust setae, merus with slender setae, carpus and propodus with robust and slender setae; dactylus with plumose seta on anteroproximal margin, with one robust and two slender setae at the base of nail. Pereopod 5 (Fig. 10C): coxa bilobed, ventral margin of posterior lobe with two robust setae, posterior margin of posterior lobe with five short setae; basis broad, subrectangular, posterior margin smooth, with numerous short setae; anterior and posterior margins of carpus with robust setae; propodus with robust setae on anterior margin, with one cluster setae on posteromedial submargin; dactylus with plumose seta on posteroproximal margin, with one robust and two slender setae at the base of nail. Pereopod 6 (Fig. 10D): coxa bilobed, anterior margin of anterior lobe with slender setae, ventral margin of posterior lobe with three robust setae, posterior margin of posterior lobe with five short setae; basis weakly expanded proximally, tapering distally, posterior margin smooth with numerous short setae; anterior margin of carpus with robust and slender setae, posterior margin with robust setae; anterior margin of propodus with robust setae, with two clusters setae on posteromedial submargin; dactylus with plumose seta on posteroproximal margin, with one robust seta and two slender setae at the base of nail. Pereopod 7 (Fig. 10E): coxa rounded, with slender setae on anterior and short setae on posterior margins; basis expanded, subovate, posterior margin rounded, smooth, with numerous short setae; anterior and posterior margins of carpus with robust and slender setae; anterior margin of propodus with robust and slender setae, with three clusters setae on posteromedial submargin; dactylus with plumose seta on posteroproximal margin, with one robust and two slender setae at the base of nail.

Figure 10. 

Elasmopus lumbiniger sp. nov. holotype male, 9.8 mm (NSMT-Cr 33064). A–E. Left pereopod 3–7, lateral view, coxal gills omitted; F. Left pleopod 1, posterior view; G. Retinacula of left pleopod 1, posterior view; H. Left pleopod 2, anterior view; I. Retinacula of left pleopod 2, anterior view; J. Left pleopod 3, anterior view; K. Retinacula of left pleopod 3, anterior view. Scale bars: 0.5 mm (A–F, H, J); 0.02 mm (G, I, K).

Coxal gills on gnathopod 2 and pereopods 3–6.

Pleopods (Fig. 10F–K): peduncles with long setae, inner distal corner of peduncle with two retinacula (Fig. 10G, I, K), peduncle of pleopod 3 broader than pleopod 1 and 2.

Uropods. Uropod 1 (Fig. 11A): peduncle length 1.2 × length of outer ramus, with six dorsolateral, seven dorsomedial and one basofacial robust setae; outer ramus slightly shorter than inner ramus, medial and lateral margins each with four robust setae, distal part with four robust setae; inner ramus medial and lateral margins each with four robust setae, distal part with five robust setae. Uropod 2 (Fig. 11B): peduncle length 0.9 × length of outer ramus; outer ramus slightly shorter than inner ramus, with three and five robust setae on medial and lateral margins, respectively, distal part with four robust setae; inner ramus five and four robust setae on medial and lateral margins, respectively, distal part with five robust setae. Uropod 3 (Fig. 11C): peduncle length 0.8 × length of outer ramus, with pair of robust setae on outer margin; rami distally truncated; outer ramus length 1.5 × length of inner ramus, with three and one clusters of robust setae on lateral and distal margins, respectively, with robust seta and one pair of robust setae on medial margin and surface, respectively; inner ramus with cluster of robust setae distally and two robust setae on medial margin.

Figure 11. 

Elasmopus lumbiniger sp. nov. A–D. Holotype male, 9.8 mm (NSMT-Cr 33064); E–I. Paratype female, 8.7 mm (NSMT-Cr 33068). A. Left uropod 1, lateral view; B. Left uropod 2, lateral view; C. Right uropod 3, medial view; D. Telson, dorsal view; E. Right gnathopod 1, medial view; F. Palmar margin of right gnathopod 1 propodus, medial view; G. Left gnathopod2, lateral view, coxal gill and oostegite omitted; H. Palmar margin of left gnathopod 2 propodus, lateral view; I. Left pereopod 3, lateral view. Scale bars: 0.3 mm (A–C); 0.1 mm (D, F, H); 0.5 mm (E, G, I).

Telson (Fig. 11D): broader than long, cleft for 75% of length; lobe trapezoid, with shallow incision at apex, each lobe bearing one plumose five robust setae apically, lateral margin of each lobe with one or two plumose setae.

Description of female

(paratype, NSMT-Cr 33068), sexually dimorphic characters. Gnathopod 1 (Fig. 11E, F): propodus anterior margin with 1 single and 4 clusters of setae, medial surface with 6 clusters of setae. Gnathopod 2 (Fig. 11G, H): slender setae on carpus anterodistal corner reaching width of carpus; propodus with two clusters, three pairs, and two2 single setae on anterolateral submargin, length of these setae 0.7 × width of propodus, anteromedial submargin with 13 clusters; palmar margin (Fig. 11H) almost straight, with minutely serrate on the distal 1/5 of the palmar margin. Pereopod 3 (Fig. 11I): basis with slender setae on anterior margin, the longest seta 0.8 × width of basis. Oostegites on gnathopod 2 and pereopods 3–5, slender with long setae.

Variations.

Mandible : left and right incisors with two or three and three or four teeth, respectively; lacinia mobilis with four or five teeth. Gnathopod 2: length of setae on carpus anterodistal corner 0.8–1.1 and 0.9–1.1 × width of carpus in male and female, length of setae on propodus anterolateral submargin 0.3–0.7 and 0.6–0.7 × width of propodus in male and female, respectively. Telson: each lobe bearing 4–6 robust setae apically.

Large male > 10 mm bearing slender setae on ventral margins of epimeral plates 2 and 3, and urosomite 1 (Fig. 12).

Figure 12. 

Elasmopus lumbiniger sp. nov., paratype male, 13.8 mm (NSMT-Cr 33073). Abdomen, lateral view. Scale bar: 0.5 mm.

Coloration in life.

Male: eyes black; antenna 1 peduncular articles brown, flagellum with two patterns, brown proximally and black distally, on a white background; body color varies among individuals ranging from reddish brown to brown or black, with small white dot patterns on head, pereon, abdomen, and coxae, with one dark black band along dorsal midline of pleonite 3 and urosomite 1. Female: body black, without distinct black band on dorsal midline of abdomen.

Distribution.

Japan: Chiba, Kanagawa, Wakayama, and Tottori (present study) (Fig. 1).

Ecological note.

This new species was found among brown and red algae or under stones in rocky shore. E. lumbiniger sp. nov. and E. projectus were collected from the same individual algae holdfasts at two sites: Kannonzaki Coast (Fig. 1A) and Uradome Beach (Fig. 1H).

Etymology.

The specific name lumbiniger, from the Latin lumbus (= loin, lower back) and niger (= black), refers to the dark coloration along the dorsal midline of male pleonite 3 and urosomite 1 in life (Fig. 6A). The Japanese name is derived from this band, which resembles a vein running along the back of a shrimp, known as “Sewata”.

Remarks.

Elasmopus lumbiniger sp. nov. is morphologically similar to E. antennatus (Stout, 1913), E. balkomanus Thomas & Barnard, 1988, and E. lemaitrei Ortiz & Lalana, 1994, in having the following features: 1) epimeral plate 3 posteroventral corner produced into well-developed acute spine; 2) male gnathopod 2 propodus palm with dense slender setae; 3) male gnathopod 2 mid-palmar margin with ridge; 4) male gnathopod 2 propodus with hump on near insertion of dactylus; 5) pereopods 5–7 basis posterior margin smooth, without long setae; and 6) telson broader than long. However, E. lumbiniger sp. nov. can be distinguished from the above three species by length of the longest seta on the anterodistal corner of male gnathopod 2 carpus 0.8–1.1 × width of carpus (less than 0.2 in these three species). From E. antennatus, this new species is distinguishable by the following two characters in addition (features of E. antennatus in parentheses): 1) hump on propodus of male gnathopod 2 trapezoidal, produced (semioval, weekly produced); and 2) the length of setae on anterior margin of pereopods 5–7 merus exceeding half the width of each merus (less than half of the width of each merus). Elasmopus lumbiniger sp. nov. can be further distinguished from E. balkomanus and E. lemaitrei by the following two additional features (features of E. balkomanus and E. lemaitrei in parentheses): 1) ridge on mid-palmar margin of male gnathopod 2 propodus not serrate (serrate); and 2) the length of setae on posterior margin of pereopod 7 carpus almost equal to that on anterior margin (more than twice that on anterior margin).

In this study, we calculated the genetic distances of COI among five Elasmopus species recorded in East Asia and Elasmopus lumbiniger sp. nov. E. lumbiniger sp. nov. differs from the other five Elasmopus species by large genetic distances (0.158–0.205 in p-distance; 0.181–0.244 in K2P), which were larger than the intraspecific distances among the Elasmopus species examined in this study (Tables 3, 4). In addition, E. lumbiniger sp. nov. is morphologically distinguished from its congeners. Therefore, it can be concluded that E. lumbiniger sp. nov. is a distinct new species.

Elasmopus mukuinu Sir & White, 2022

Figs 13, 14, 15 [New Japanese name: Mukuinu-iso-yokoebi]

Elasmopus mukuinu Sir & White, 2022: 574, figs 1C, D, 4–9.

Material examined.

• Male, 9.3 mm (NSMT-Cr 33079: Fig. 13A), Kaichu Doro, Uruma, Okinawa, Japan (26.3324°N, 127.9222°E) (Fig. 1J), intertidal zone, coral beach, among the green alga Ulva sp., 14 March 2025, H. Yoshimura coll.; • ovigerous female, 7.5 mm (NSMT-Cr 33080), Ono Beach, Shimoda, Shizuoka, Japan (34.6329°N, 138.8975°E) (Fig. 1E), shallow subtidal zone, rocky shore, under the stone, 24 February 2024, H. Yoshimura coll.; • male, 8.7, 7.4 mm (NSMT-Cr 33081: Fig. 13B, NSMT-Cr 33082), ovigerous female, 5.1 mm (NSMT-Cr 33083: Fig. 13C), Oura Beach, Shikinejima, Tokyo, Japan (34.3307°N, 139.2083°E) (Fig. 1F), shallow subtidal zone, rocky shore, under the stone, 3 August 2024, H. Yoshimura coll.; • male, 14.6, 7.9, 6.5 mm (NSMT-Cr 33084–33086), Yakengahama Beach, Hachijojima, Tokyo, Japan (33.1007°N, 139.7696°E) (Fig. 1G), rocky shore, 20 December 2022, K. Tomikawa coll.

Figure 13. 

Elasmopus mukuinu. A. Male, 9.3 mm (NSMT-Cr 33079), lateral view; B. Male, 8.7 mm (NSMT-Cr 33081), lateral view; C. Female, 5.1 mm (NSMT-Cr 33083), lateral view. Scale bars: 1.0 mm.

Type locality.

Kaichu Doro, Okinawa Island, Japan (26.3321°N, 127.9146°E).

Diagnosis.

Epimeral plate 3 posterior margin weakly serrated. Antenna 1 peduncles bearing slender setae on lateral and medial margins, with bi- or tri-articulate accessory flagellum. Mandibular palp article 3 short. Gnathopod 2 carpus anterodistal corner with slender setae, length of the longest seta reaching 0.3–0.6 and 0.4–0.6 × width of carpus in male and female. Male gnathopod 2 propodus posterior margin and inner surface with dense slender setae, length of the longest seta 0.9 × width of propodus, palmar margin with semicircular hump near hinge of dactylus, mid-palmer margin with ridge. Pereopods 5–7 basis posterior margin smooth, without long setae. Telson cleft, border than long; lobe trapezoid, with shallow incision at apex, inner apical corner rounded and produced, each lobe bearing one plumose and one or two robust setae apically.

Description of male

(NSMT-Cr 33081). Body (Fig. 13B): smooth, not carinate, with a few short setae dorsally on pereon and abdomen. Head: eyes oval; lateral cephalic lobe broad, anteroventral margin with notch; dorsal surface with a few short setae. Epimeral plate 3 (Fig. 14A): ventral margin with two pairs and four single robust setae, posterior margin weakly serrated, with four short setae.

Figure 14. 

Elasmopus mukuinu. A–F. Male, 8.7 mm (NSMT-Cr 33081); G. Female, 5.1 mm (NSMT-Cr 33083). A. Right epimeral plate 3, lateral view; B. Right accessory flagellum, medial view; C, D. Incisor, lacinia mobilis, accessory setal row and molar process of left and right mandible, medial view; E. Right mandibular palp articles 2, 3, medial view; F. Telson, dorsal view; G. Right accessory flagellum, medial view. Scale bars: 0.3 mm (A); 0.1 mm (B–G).

Antenna 1 : length 0.6 × body length; peduncular articles 1–3 with slender setae on lateral and medial margins; accessory flagellum (Fig. 14B) tri-articulate, reaching to distal margin of primary flagellar article 1, accessory flagellar article 3 tiny; primary flagellum 28-articulate, with slender setae.

Mandible (Fig. 14C–E) with left and right incisors three and four teeth, respectively; left and right lacinia mobilis three and four teeth, respectively; accessory setal row consisting of four setae on each of left and right mandibles; molar process well developed, triturative; palp well developed, tri-articulate; article 2 with seven setae, article 3 falcate, length 3.2 × width.

Gnathopod 1 (Fig. 15A): subchelate; coxa anteroventral corner weakly produced; basis with long setae on posterior margin and medial surface; carpus subequal in length to propodus, with slender seta on anterodistal corner, with dense setae on posterior margin and medial surface; propodus with four and five clusters of setae on anterior margin and medial surface, respectively, posterior margin with row of slender and robust setae, palmar margin almost transverse, minutely serrate, with rows of robust setae on medial and lateral palmar submargins. Gnathopod 2 (Fig. 15B): subchelate; basis with long setae on posterior margin; carpus with slender setae on anterodistal corner, length of these setae reaching 0.6 × width of carpus, with dense setae on posterior margin; propodus with one cluster, one pair and three single setae on anterolateral submargin, length of these setae reaching 0.2 × width of propodus, with nine clusters of setae on anteromedial submargin, lateral palmar margin with hump on near insertion of dactylus, mid-palmar margin with ridge, posterior margin with row of plumose setae, the proximal seta longer than the distal seta, length of the longest seta reaching 0.9 × width of propodus.

Figure 15. 

Elasmopus mukuinu. A, B. Male, 8.7 mm (NSMT-Cr 33081); C, D. Female, 5.1 mm (NSMT-Cr 33083). A. Right gnathopod 1, medial view; B. Right gnathopod 2, medial view; C. Right gnathopod 1, medial view; D. Right gnathopod 2, medial view. Scale bars: 0.3 mm.

Telson (Fig. 13F): broader than long, cleft for 65% of length; lobe trapezoid, with shallow incision at apex, inner apical corner rounded and produced, each lobe bearing one plumose and two robust setae apically, lateral margin of each lobe with one plumose seta.

Description of female

(NSMT-Cr 33083), sexually dimorphic characters. Antenna 1: accessory flagellum (Fig. 14G) bi-articulate. Gnathopod 1 (Fig. 15C): propodus anterior margin with three clusters of setae, medial surface with five clusters of setae. Gnathopod 2 (Fig. 15D): slender setae on carpus anterodistal corner 0.6 × width of carpus; propodus with four slender setae on anterolateral submargin, the longest setae 0.4 × width of propodus, with eight and ten clusters of setae on anteromedial margin and medial surface, respectively; palmar margin almost straight, with minutely serrate on the distal 2/5 of the palmar margin.

Variations.

Mandible : both incisors with 2–4 teeth; right lacinia mobilis with four or five teeth. Gnathopod 2: length of setae on carpus anterodistal corner 0.3–0.6 and 0.4–0.6 × width of carpus in male and female, length of setae on propodus anterolateral submargin 0.2–0.3 and 0.3–0.5 × width of propodus in male and female, respectively. Telson: each lobe bearing one or two robust setae apically.

Coloration in life.

Eyes black; antennae 1 and 2 peduncular articles brown, flagellum articles have two black patterns on a white background; body translucent white, laterally brown, with small white dot patterns on head, pereon, and abdomen, with orange pattern on epimeral plates. Male and female have similar body coloration.

Distribution.

Japan: Shizuoka, Izu Islands (Shikinejima Island and Hachijojima Island), Tokyo (present study); Okinawa Island, Okinawa (Sir and White 2022; present study) (Fig. 1).

Remarks.

The examined specimens agreed well with the original description of E. mukuinu by Sir and White (2022), but there was regional morphological variation in the number of accessory flagellum articles of antenna 1. In the male from Okinawa Island (Fig. 1J) and females from Shizuoka (Fig. 1E) and Shikinejima Island (Fig. 1F), the accessory flagellum consisted of two articles, consistent with the original description (like Fig. 14G). In contrast, males from Shikinejima Island and Hachijojima Island (Fig. 1G) possessed a tri-articulate accessory flagellum (like Fig. 14B). Therefore, the number of accessory flagellum articles in males differed between localities: E. mukuinu from Okinawa Island had two articles, whereas those from the Izu Islands (Shikinejima and Hachijojima Islands) had three.

The genetic distance of COI between E. mukuinu from Okinawa Island and those from Shizuoka and the Izu Islands was 0.050 (p-distance) and 0.052 (K2P), respectively, representing the largest intraspecific genetic distance among the Elasmopus species examined in this study (Tables 3, 4). However, we believed that these morphological and genetic differences were not sufficient to support separation into distinct species, and we concluded that these specimens should be regarded as conspecific. Further examination of E. mukuinu specimens from other regions will help clarify the taxonomic status of local populations of this species.

Genetic distances among Elasmopus species

The intra- and interspecific distances in Elasmopus were calculated using p-distance (Table 3) and K2P distance (Table 4). The maximum intraspecific distances were 0.050 (p-distance) and 0.052 (K2P), and the minimum interspecific distances were 0.158 (p-distance) and 0.181 (K2P).

Key to the species of Elasmopus from East Asia

The following dichotomous key excludes E. spinimanus, whose occurrence in East Asia is doubtful.

1 Urosomite 1 with triangular process dorsally E. japonicus
Urosomite 1 dorsally smooth 2
2 Male gnathopod 2 propodus medial surface and palmar margin with few setae 3
Male gnathopod 2 propodus medial surface and palmar margin with dense slender setae 6
3 Antenna 2 flagellum articles anteroposteriorly compressed; pereopods 5–7 bases without long setae on posterior margins E. koreanus
Antenna 2 flagellum articles nearly cylindrical; pereopods 5–7 bases with long setae on posterior margins 4
4 Telson shorter than wide E. hooheno
Telson longer than wide 5
5 Foremost medial tooth on palm of male gnathopod 2 not exceeding palmar margin; posterodistal setae on propodi of pereopods 5–7 longer than length of dactyli E. rapax
Foremost medial tooth on palm of male gnathopod 2 exceeding palmar margin; posterodistal setae on propodi of pereopods 5–7 shorter than length of dactyli E. nkjaf
6 Posterior margin of pereopod 6 basis castelloserrate 7
Posterior margin of pereopod 6 basis smooth 8
7 Posterior margin of pereopod 7 basis castelloserrate; epimeral plate 3 posteroventral corner with distinct spine E. nanshaensis
Posterior margin of pereopod 7 basis smooth; epimeral plate 3 posteroventral corner notched, without distinct spine E. alkhiranensis
8 Inner margins of pereopods 3–7 dactyli serrate E. spinidactylus
Inner margins of pereopods 3–7 dactyli smooth 9
9 Male gnathopod 2 carpus with a row of slender setae along anterior margin E. spinicarpus
Male gnathopod 2 carpus anterior margin with setae on anterodistal corner only, or on both anterodistal corner and mid-anterior margin 10
10 Anterior setae on meri of pereopods 5–7 shorter than half width of meri E. hawaiensis sensu Ren, 2012
Anterior setae on meri of pereopods 5–7 longer than half width of meri 11
11 Male gnathopod 2 propodus mid-palmer margin with well-developed process; length of setae on gnathopod 2 carpus anterodistal corner less than 0.2 × width of carpus in male, less than 0.4 × in female; male pereopod 7 merus to propodus without slender setae on posterior margins E. projectus
Male gnathopod 2 propodus mid-palmer margin with ridge; length of setae on gnathopod 2 carpus anterodistal corner exceeding 0.3 × width of carpus in male, exceeding 0.4 × in female; male pereopod 7 merus to propodus with slender setae on posterior margins 12
12 Epimeral plate 3 posteroventral corner without distinct projection; length of setae on gnathopod 2 carpus anterodistal corner 0.3–0.6 and 0.4–0.6 × width of carpus in male and female; telson each lobe bearing 1 or 2 robust setae apically E. mukuinu
Epimeral plate 3 posteroventral corner produced into well-developed acute spine; length of setae on gnathopod 2 carpus anterodistal corner 0.8–1.1 and 0.9–1.1 × width of carpus in male and female; telson each lobe bearing 5 or 6 robust setae apically E. lumbiniger sp. nov.

Discussion

Species diversity of Elasmopus in East Asia

To date, thirteen species of Elasmopus have been recorded in East Asia. In the southern limit of this region, eight species are known from South China Sea (Table 1; Ren 2012; Myers and Momtazi 2015): E. alkhiranensis, E. hawaiensis, E. hooheno, E. nanshaensis, E. rapax, E. spinicarpus, E. spinidactylus, and E. spinimanus. Among these, the record of E. spinimanus by Ren (2012) was based solely on a single female specimen. This species was originally described from Sri Lanka based on mainly male specimens (Walker 1904). The female morphological characters of E. spinimanus were later described from specimens collected in Fiji (Myers 1985). We compared the female morphology of E. spinimanus described by Ren (2012) and Myers (1985) and found that they closely agreed. However, accurate species identification within Elasmopus requires examination of male morphological futures. Therefore, the identification of E. spinimanus by Ren (2012) is doubtful. Further taxonomic investigation, including examination of male specimens from the same population of E. spinimanus sensu Ren (2012), is necessary to confirm its taxonomic position.

At its northern limit, E. smirnovi Bulyčeva, 1952 was described from Primorsky Krai, Russian Far East (Bulyčeva 1952). However, this species exhibits morphological characteristics that differ from those of Elasmopus and is suggested to belong to the Maera group (Vader and Krapp-Schickel 2012). Because E. smirnovi could not belong to Elasmopus, Japanese and South Korean waters represent the true northernmost distribution of this genus in the northwestern Pacific and are important for understanding its diversity. Until now, only two species, E. japonicus and E. projectus, had been recorded in temperate regions of Japan, and two species, E. koreanus and E. rapax, from South Korea. However, the present study recorded three additional species from temperate Japan: E. koreanus, E. lumbiniger sp. nov., and E. mukuinu. These findings suggest that the species diversity of Elasmopus in northern East Asia is higher than previously recognized. Further surveys in previously unstudied regions of Japan and South Korea could find additional unrecorded or undescribed species.

Conclusions

Elasmopus lumbiniger sp. nov., found in Japan, is distinguished from its congeners by its morphological features. Elasmopus koreanus, previously known only in Korea, is newly recorded in Japan. Elasmopus mukuinu, which has been reported only from Okinawa Island, is also found along the temperate coast of Japan, greatly extending its known distribution range. These findings shed light on the high species diversity of Elasmopus in waters around Japan and South Korea. Furthermore, the nucleotide sequences of the COI of these species are provided.

Acknowledgements

We thank Yukiko Mukaida for her critical reading of the manuscript and support in collecting specimens. We are also grateful to Hiroshi Ogawa and Masako Ooga for their assistance with the field surveys. Our sincere thanks go to Alan A. Myers and Jesser Fidelis de Souza Filho for their constructive and valuable comments on this work. We would also thank Luiz F. Andrade for revising an earlier version of the manuscript. We extend our appreciation to the Natural Science Center for Basic Research and Development for granting access to experimental facilities.

Additional information

Conflict of interest

The authors have declared that no competing interests exist.

Ethical statement

No ethical statement was reported.

Use of AI

No use of AI was reported.

Funding

This work was partly supported by the Japan Society for the Promotion of Science (JSPS) through Grant-in-Aid for JSPS Fellows [JP25KJ1881] to H. Yoshimura, and through KAKENHI [JP22H01011 and JP22K06373] to K. Tomikawa. Additional support was provided by a grant from the Nakdonggang National Institute of Biological Resources (NNIBR), funded by the Ministry of Environment (MOE) of the Republic of Korea [NNIBR20252102] to C.W. Lee.

Author contributions

Writing - original draft: HY. Writing - review and editing: KT, CWL.

Author ORCIDs

Hiro Yoshimura https://orcid.org/0009-0005-1297-4368

Chi-Woo Lee https://orcid.org/0000-0002-0763-4271

Ko Tomikawa https://orcid.org/0000-0003-1521-9016

Data availability

All of the data that support the findings of this study are available in the main text or Supplementary Information.

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