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Research Article
A new species of Nebalia (Crustacea, Leptostraca) from a hydrothermal field in Kagoshima Bay, Japan
expand article infoTakuma Hirata, Yoshihiro Fujiwara§, Tomohiko Kikuchi
‡ Yokohama National University, Yokohama, Japan
§ Japan Agency for Marine-Earth Science and Technology, Yokohama, Japan
Open Access

Abstract

A new species of Leptostraca, Nebalia tagiri sp. nov. is described and illustrated. This species was sampled from 200 m depth at a hydrothermal field in Wakamiko Caldera of Kagoshima Bay, Japan. Nebalia tagiri sp. nov. is different from known Nebalia species as follows: rostral length 2.4 times as long as width; article 4 of antennule with 3–5 robust distal spines; antennular scale approximately twice as long as wide; article 3 of antenna with eight spines and nine spine-like setae along proximal half, two thin setae and six spine-like setae on external lateral face, six spines and four simple setae on distal margin; article 1 of second maxilla longer than article 2; article 2 of mandibular palp with two thin setae; exopod of pleopod 1 with 21 spines along lateral margin; furcal rami longer than combined length of pleonite 7 and telson; rounded denticles of pleonite 6 and 7; anal-plates ‘shoulder’ not distinct. Furthermore, this specimen is the first genus Nebalia found in the hydrothermal vent. The distribution and ecology of this new species is also discussed and a key to all species of Nebalia is provided.

Keywords

Hydrothermal vent, key, Leptostraca, Malacostraca, Nebalia, new species

Introduction

The genus Nebalia is a member of the Order Leptostraca, Suborder Nebaliacea, Family Nebaliidae. The family includes four other genera: Dahlella (Hessler, 1984), Nebaliella (Thiele, 1904), Sarsinebalia (Dahl, 1985), and Speonebalia (Bowman et al., 1985). Nebalia was established by Leach (1814) with type Nebalia herbstii from the British Isles. More than 37 species of this genus have been reported thus far from Africa (Barnard 1914; Kensley 1976; Olesen 1999; Bochert and Zettler 2012), Red Sea (Wägele 1983), Adriatic Sea (Dahl 1985), Britain-Celtic Sea (Dahl 1985), Greenland (Dahl 1985), Norway (Dahl 1985), Pakistan (Kazmi and Tirmizi 1989), Antarctic Sea (Dahl 1990), Falkland Islands (Dahl 1990), New Zealand (Dahl 1990), South Atlantic Ocean (Dahl 1990), Mexico (Escobar-Briones and Villalobos-Hiriart 1995; Ortiz et al. 2011), California (Martin et al. 1996; Vetter 1996; Haney and Martin 2000, 2005), Mediterranean Sea (Ledoyer 1997; Moreira et al. 2007, 2012; Koçak and Moreira 2015), New Caledonia (Ledoyer 2000), Northeast Atlantic (Haney et al. 2001; Moreira et al. 2003, 2009), Aegean Sea (Moreira et al. 2007), Hong Kong (Lee and Bamber 2011), South Korea (Song et al. 2012, 2013; Song and Min 2017), and Malaysia (Othman et al. 2016). Although our understanding of the morphology and distribution of the genus Nebalia has progressed, further investigation into the ecology of the genus is necessary.

In 2008, a survey was undertaken at the hydrothermal field of the Wakamiko Caldera in Kagoshima Bay, Japan using the deep-sea remotely operated vehicle (ROV) “Hyper-Dolphin” of JAMSTEC. Twenty-eight specimens of a previously undocumented species in the genus Nebalia were discovered. In this paper, we describe these specimens as Nebalia tagiri sp. nov. Taxonomic keys used to identify all currently known Nebalia species are also provided. Comments relating to the ecology and distribution of this species are also noted.

Materials and methods

Samples were collected in 2008 at a hydrothermal vent in the Wakamiko Caldera, Kagoshima Bay, during dive number 886 of the Japanese deep-sea ROV “Hyper-Dolphin” of JAMSTEC. The sampling site was the Wakamiko Caldera, located north of Mt. Sakurajima of Kagoshima Bay at a depth of ca. 200 m (Fig. 1). The water temperature in this field is ca. 10 °C. While bacterial mats were observed, these hydrothermal vents have no documented epifauna occurring around them. Specimens examined in this study were captured using a suction sampler installed on the ROV. Samples were sorted, identified, sexed, and measured. Total length (TL: measured from the articulation between the rostrum and the carapace to the posterior end of the caudal furca), carapace length (CL: measured from the antero-dorsal margin of carapace to the posterio-median margin of carapace), and rostrum length (RL: measured along the midline) were used as size indicators. Illustrations were made with the aid of a camera lucida. The type materials were deposited at the National Science Museum (Natural History), Tokyo (NSMT), with the remaining material deposited in the JAMSTEC Marine Biological Sample repository.

Figure 1. 

Sampling location (in yellow) of Nebalia tagiri sp. nov. in Kagoshima Bay.

Systematics

Nebalia Leach, 1814

Diagnosis

Carapace almost reaching medial margin of pleonite 4. Rostrum long and narrow, 2.4 times as long as wide. Surface of eyes smooth. Article 4 of antennule with row of four simple setae and four robust distal spines. Article 3 of antenna with eight spines and nine spine-like setae along proximal half, two thin setae and six spine-like setae on external lateral face, six spines and four simple setae on terminal margin. Article 1 of endopod of second maxilla longer than article 2. Exopod of second maxilla longer than article 1 of endopod. Article 2 of mandibular palp with two thin setae. Pleonites 6 and 7 with distally rounded denticles along posterior border. Exopod of pleopod 1 with a single row of approximately 21 stout serrated spines along lateral margin. Anal plates with no distinct lateral ‘shoulder’. Furcal rami longer than combined length of pleonite 7 and telson.

Nebalia tagiri sp. nov.

Figs 2, 3, 4, 5, 6, 7

Material

Twenty-eight specimens were collected using the JAMSTEC ROV “Hyper-Dolphin” of JAMSTEC during dive 886 in the Wakamiko Caldera during the R/V “Natsushima” NT08-17 Leg-1 cruise. “Hyper-Dolphin” dive 886: the Tagiri Site on the Wakamiko Caldera: 30°40.068'N, 130°45.690'E; 200 m; 7 Aug 2008. 14 ♂♂ (TL: 3.7–6.0 mm), 13 ♀♀ (TL: 2.0–6.1 mm).

Types

Holotype : (NSMT-Cr 26758), adult ♀ of 10.7 mm TL. Allotype: (NSMT-Cr 26759), adult ♂ of 7.0 mm TL. Paratypes: 3 adult ♂♂ (TL: 4.1–6.0 mm) (NSMT-Cr 26760, NSMT-Cr 26761, NSMT-Cr 26762) 3 adult ♀♀ (TL: 3.9–6.0 mm) (NSMT-Cr 26763, NSMT-Cr 26764, NSMT-Cr 26765).

Description

Female holotype. Carapace (Fig. 2A) oval, ca. 1.5 times as long as wide, almost reaching pleonite 4.

Rostrum (Fig. 2B) long and narrow, 2.4 times as long as width, with round apex.

Compound eye (Fig. 2C): ommatidial part covering two-thirds of eye-stalk. Supraocular plate reaching to ommatidial part.

Antennule (Fig. 2D): peduncle composed of four articles. Article 2 longer than article 3, with single long plumose seta on anterior margin, 5 long and three short plumose setae arising subterminally and cluster of simple setae on anterior margin, respectively. Article 3 shorter than article 2, widest distally, with terminal cluster of simple setae and long simple seta arising on anterior margin and five long plumose setae and two thin plumose setae on posterio-distal margin. Article 4 much shorter than article 3, with row of four simple setae and four robust spines distally. Antennular scale oval, twice as long as width. Flagellum slightly longer than peduncle, composed of 12 articles.

Antenna (Fig. 2E): peduncle composed of 3 articles. Article 2 2.3 times as long as wide, with stout spine at dorso-distal portion. Article 3 longer than article 2, with different pattern of spines or setae along medial anterior margin as follows:

(1) proximal row of ca. six simple setae and plumose seta on inner surface;

(2) eight spines and nine spine-like setae along proximal half, the distalmost being the longest, respectively;

(3) two thin setae and six spine-like setae on external lateral face;

(4) seven thin plumose setae, three plumose setae and seven simple setae, each associated proximal spines;

(5) six robust spines increasing in length distally and four simple setae at apex, 21 long plumose setae arising from posterior distal margin, and robust plumose seta arising sub-terminally. Flagellum longer than peduncle, composed of 15 articles.

Mandible (Fig. 2F) well developed. Mandibular palp composed of three articles. Article 2 equal in length as article 3, article 2 with two thin setae at mid-length on lateral margin and sub-terminal on superior margin, respectively. Article 3 cylindrical, with marginal setae-row covering four-fifths length of article. Molar process shorter than palp article 1, distal margin with row of teeth forming grinding surface. Incisor process broad basally with acute terminal process and minute tooth along lateral margin.

First maxilla (Fig. 2G): proximal endite (Fig. 2H) with rounded medial margin, bearing short robust plumose setae. Distal endite (Fig. 2I) carrying two rows of stout, spatulate setae and two long plumose seta. Palp long, ca. 4.5 times longer than combined length of both endites, bearing 20 setae.

Second maxilla (Fig. 2J): protopod sub-divided into four endites bearing plumose setae. Endite 1 rectangular and endite 3 rounded approximately equally sized to endite 1; endite 2 oval, smaller than endite 1 and 3; endite 4 smaller than other endites. Endopod composed of two articles, article 1 ca. twice as long as article 2, lateral margin with plumose setae, article 2 with three terminal plumose setae. Exopod reaching beyond apex of endopod article 1, bearing 23 plumose setae on lateral margin.

Figure 2. 

Nebalia tagiri sp. nov. A female holotype, lateral view B rostrum C eye D antennule E antenna and detail of different row of spines and setae of article 3 F mandible and detail of incisor process G first maxilla H detail of proximal endite of first maxilla I detail of distal endite of first maxilla J second maxilla.

Thoracopod 1 (Fig. 3A): endopod composed of large article and three small distal articles, with numerous plumose setae along outer margin, terminal setae not reaching the terminal margin of exopod. Exopod oval, not reaching beyond the terminal article of endopod, with single long setae on terminal margin and 21 thin setae along inner margin. Epipod large, proximal lobe beyond the basis, distal lobe reaching beyond the middle of exopod.

Thoracopod 2–6 (Fig. 3B–F): Except for the exopod, shape of each limbs same. Eleven thin setae of inner margin of exopod in thoracopod 2, five in thoracopod 3, seven in thoracopod 4, six in thoracopod 5, seven in thoracopod 6. Shape of exopod gradually changes from oval to triangular from thoracopods 2–6.

Thoracopod 7 (Fig. 3G) endopod composed of one large article and two small distal articles, with numerous plumose setae along outer margin, terminal setae beyond the terminal margin of exopod. Exopod rounded and distal part expanded like a triangle, reaching beyond the terminal article of endopod, with eight thin setae along inner margin. Epipod large and triangular, proximal lobe beyond the basis, distal lobe reaching beyond the middle of exopod, with three thin setae along distal margin.

Thoracopod 8 (Fig. 3H): smaller than other thoracopods. Endopod composed of a large article and three small distal articles, with numerous plumose setae along outer margin, terminal setae not reaching the terminal margin of exopod. Exopod oval, reaching beyond the terminal article of endopod, with five setae along inner margin. Epipod small, proximal lobe beyond the basis, distal lobe not reaching beyond the middle of exopod.

Figure 3. 

Nebalia tagiri sp. nov. A thoracopod 1 B thoracopod 2 C thoracopod 3 D thoracopod 4 E thoracopod 5 F thoracopod 6 G thoracopod 7 H thoracopod 8.

Pleon (Fig. 2A, 5A): composed of seven segments. Posterior margin of pleonite 1 smooth. Pleonite 2 with narrowly triangular denticles on half-length of posterior margin with wide flat margin. Pleonite 3 with round minute denticles. Pleonite 4 with round minute denticles, posterolateral margin expanded and forming narrow acute process. Pleonites 5, 6, and 7 with minute rounded denticles.

Pleopod 1 (Fig. 4A): protopod twice as long as width, with three short simple setae arising proximally, simple seta medially, two spine-like setae near base of endopod, spine-like seta near base of exopod. Endopod composed of two segments, longer than exopod, distal segment with acute process at apex, bearing long robust simple spine, lateral and medial margin each with plumose setae, nine short setae on proximal medial margin, appendix interna of proximal segment with three short recurved hooks. Exopod with row of 21 stout serrated spines along lateral margin, five stout simple spines on distolateral margin, distal one longest, plumose setae along distal inner margin.

Pleopod 2 (Fig. 4B): protopod 2.2 times as long as wide, six simple setae and seven simple setae on proximal and distal part of inner side, respectively; simple seta near base of exopod, with blade-like process between exopod and endopod. Endopod composed of two segments, longer than exopod, distal segment with acute process at apex, bearing long robust simple spine, lateral and medial margins each with plumose setae, ten short setae on proximal medial margin, appendix interna of proximal segment with three short recurved hooks. Exopod with row of six pairs of robust spines along lateral margin, three stout simple spines on distal margin, plumose setae along distal inner margin.

Pleopod 3 (Fig. 4C): protopod 2.2 times as long as wide, five simple setae on proximal and distal part of inner side respectively, simple seta near base of exopod, with blade-like process between exopod and endopod. Endopod composed of two segments, longer than exopod, distal segment with acute process at apex, bearing long robust simple spine, lateral and medial margin each with plumose setae, eight short setae on proximal medial margin, appendix interna of proximal segment with three short recurved hooks. Exopod with row of seven pairs of robust spines along lateral margin, three stout simple spines on distal margin, plumose setae along distal inner margin.

Pleopod 4 (Fig. 4D): protopod twice as long as width, bearing five simple setae along lateral proximal border, eleven short simple setae along ventral proximal border and four simple setae along posterior border. Posterolateral corner of peduncle produced as acute point, with blade-like process between exopod and endopod, posterior margin lacking serrations, posterolateral corner with acute projection. Endopod composed of two segments, longer than exopod, distal segment of endopod with acute process at apex, bearing long robust simple spine, lateral and medial margin each with plumose setae, ten short setae on proximal medial margin, appendix interna of proximal segment with three short recurved hooks. Exopod with row of seven pairs of robust spines along lateral margin, three stout simple spines on distal margin, plumose setae along distal inner margin.

Pleopod 5 (Fig. 4E, F): acute process between bases of rami. Protopod composed of two segments. Distal segment 4.5 times as long as wide, bearing five simple spines and ten short setae, lateral margin with ca. 33 simple setae.

Pleopod 6 (Fig. 4G, H): acute process between bases of rami, rami bearing five simple spines and six simple setae.

Figure 4. 

Nebalia tagiri sp. nov. A pleopod 1 B pleopod 2 C pleopod 3 D pleopod 4 and detail of lateral margin E pleopod 5 F detail of pleopod 5 G pleopod 6 H detail of pleopod 6.

Telson, anal plates, and furca (Fig. 5B, C): anal plates (Fig. 5C) with medial margin slightly convex, point acute, lateral margin with no distinct ‘shoulder’. Furcal rami (Fig. 5B) slightly longer than combined lengths of pleonite 7 and telson, 21 spines along outer edge, 14 robust setae and 16 plumose setae along inner margin, three long robust setae and thin spine on distolateral margin.

Figure 5. 

Nebalia tagiri sp. nov. A female pleonite, dorsal view and detail of denticles B furcal rami (setae not illustrated for left limb) C anal plates.

Color in life (Fig. 7): living specimens with dark red eyes and most of body transparent.

Allotype

(adult male): antennule flagellum more swollen proximally than in female (Fig. 6B). Antenna flagellum composed of more than 50 articles (Fig. 6C). article 2 of endopod palp half-length of article 1 (Fig. 6D). Furcal rami sub-equal in length to combined length of pleonite 7 and telson (Fig. 6E).

Figure 6. 

Nebalia tagiri sp. nov. A male allotype, lateral view B antennule C antenna D second maxilla E furcal rami (setae not illustrated for left limb) F anal plates.

Figure 7. 

Nebalia tagiri sp. nov., photographs A female, lateral view B female with larvae C female with eggs.

Morphological variations

Examination of five female specimens of various sizes shows several morphological variations in the number of robust distal spines of article 4 of antennule and lateral spines of exopod of pleopod 1. Article 4 of antennule with 3–5 spines, exopod of pleopod 1 with 21–24 spines on lateral margin, respectively.

Etymology

The specific name tagiri originates from the Japanese word ‘tagiru’ meaning “boiling”, a reflection of the hydrothermal venting and bubbling of methane and carbon dioxide as found in the habitat of N. tagiri sp. nov.

Remarks

Nebalia tagiri sp. nov. differs from the other 34 described species based on four unique characteristics: (1) smooth oval eye-stalk; (2) ommatidial part covering two-thirds of eye-stalk; (3) article 4 of antennule with 3–5 robust distal spines; (4) article 1 of second maxilla endopod longer than article 2. Nebalia tagiri sp. nov. can easily be distinguished from N. bipes (Fabricius, 1780), N. mortoni (Lee & Bamber, 2011), and N. koreana (Song et al., 2012) based on the following: (1) length of rostrum in N. tagiri, N. koreana, and N. mortoni ca. 2.4 times as long as width, N. bipes approximately twice times as long as width; (2) number of distal spines of article 4 of antennule is four in N. tagiri sp. nov. and N. mortoni, three in N. bipes and five in N. koreana; (3) length of antennular scale in N. tagiri, N. koreana and N. bipes ca. twice as long as width, N. mortoni ca. 2.7 times as long as width; (4) the different pattern of spines or setae of article 3 of antenna among related three species are summarized in Table. 1.

Table 1.

Comparison of Nebalia tagiri sp. nov. with related species of Nebalia. Key: ro = Rostrum; an1 = antennule; an2 = antenna; mp = mandibular palp; pp = pleopod; pn = pleonite; a = article; exp = exopod; sp = spine; se = seta; sls = spine-like seta; ts = thin seta; ps = plumose seta.

Habitat Depth Ro width Shape of eye-stalk Distal sp of an1 a4 Proximalrow of an2 a3 Lateral row of an2 a3 Distal row of an2 a3 Ts of mp2 a2 Sp on pp1 exp Uropod length Shape of pn6–7 denticles Presence of ‘shoulder’ on anal–plates Reference
N. tagiri sp. nov. Hydrothermal vent chimney 200 m 2.4 times Oval 4 sp 8 sp 2 ts 6 sp 2 ts 21–24 sp > Pn7+t Round No distinct This paper
9 sls 6 sls 4 se
N. abyssicola Mud 680–820 m Oval 1 sp 8 sp 2 ts 5 sp 2 ts 25–30 sp > Pn7+t Round None Ledoyer (1997)
6 se 6 sp 6 se
3 sls
N. bipes Clay and stones 5–13 m 2 times Oval 3 sp 1 ts 30 sp = Pn7+t Round Distinct Dahl (1985)
N. borealis Sand 240 m 2.1 times Oval 2 sp 1 ts 24 sp ≥ Pn7+t Round to acute Distinct Dahl (1985)
N. koreana Algal mat 6 m 2.4 times Oval 5 sp 10 sp 2 ts 5 sp 2 ts 30–38 sp < Pn7+t Round No distinct Song et al. (2012)
6 sls 6 sls 4 se
N. mortoni 17 m 2.4 times Oval 4 sp 7 sp 2 ts 6 sp 2 ts 25 sp = Pn7+t Square No distinct Lee and Bamber (2011)
4 sls
N. schizophthalma 2886 m 2.6 times Bilobed 5 ts 10 sls 10 se 6 sp 2 ts 15 sp < Pn7+t Acute None Haney et al. (2001)
8 se

Nebalia tagiri sp. nov. showed different characteristics from N. koreana or N. mortoni in the following points: (1) number of spines or spine-like setae along proximal half in comparison with N. koreana and N. mortoni; (2) existence of spine-like setae on external lateral face in comparison with N. mortoni; (3) number of spines and setae on terminal margin in comparison with N. koreana and N. mortoni, N. bipes is lacking detailed description; (5) thin seta of article 2 of mandibular palp is two in N. tagiri sp. nov., N. koreana and N. mortoni and one in N. bipes; (6) number of lateral spines of exopod of pleopod 1 is 21–24 in N. tagiri sp. nov, 25 in N. mortoni, and more than 30 in N. bipes and N. koreana; (7) furcal rami are longer than the combined length of pleonite 7 and the telson in N. tagiri. sp. nov. and of the same length in N. bipes and N. mortoni, while in N. koreana they are shorter; (8) denticles of posterior margin of pleonites 6 and 7 are rounded in N. tagiri sp. nov., N. bipes, and N. koreana while in N. mortoni are square-shaped.

Most species of Nebalia have been reported from shallow water (< 10 m depth) in the world oceans with two exceptions, i.e., Nebalia abyssicola (Ledoyer, 1997) and N. schizophthlma (Haney et al., 2001) have been reported from ca. 100 m or deeper (Table 1). Nebalia schizophthalma was reported at a depth of 2886 m in the North Atlantic Ocean, which is the deepest record of this genus. Nebalia tagiri sp. nov. was collected from a hydrothermal vent chimney at a depth of 200 m in Wakamiko Caldera of Kagoshima Bay, Japan (Fig. 1). This depth of 200 m is relatively deep compared to similar species of this genus. The genus Nebalia has not been previously reported from any hydrothermal fields to date. The only species reported from a deep-sea hydrothermal field is Dahlella caldariensis (Hessler, 1984), observed on the mussel beds of hydrothermal vent areas at the Galapagos and the East Pacific Rise at depths deeper than 2000 m. On the other hand, N. tagiri sp. nov. was collected near the interior of chimney walls, which were close to the chimney vents spouting thermal water (ca. 200 °C).

This species was clearly observed on the inner surface of the chimney, suggesting the species is adapted to the hydrothermal environment. For a more accurate understanding of this species, further investigations are needed to determine their ecological and/or physiological aspects in relation to the hydrothermal fields.

Key to species of genus Nebalia

1 Pleopod 6 composed of two segments N. biarticulata Ledoyer, 1997
Pleopod 6 composed of one segment 2
2 Entire surface of eye smooth 13
Eye surface different 3
3 Dorsal margin of eye smooth, with lobes only on antero-distal margin 10
Eye papillae present on dorsal margin 4
4 Eye with several lobes on antero-distal margin N. cambodiana Song et al., 2013
Eye with antero-distal margin smooth, lacking lobes 5
5 Ommatidial part covering most of the eye-stalk 6
Ommatidial part not covering most of the eye-stalk 7
6 Protopod of pleopod 4 with serrations along posterior margin N. cannoni Dahl, 1990
Protopod of pleopod 4 lacking serrations along posterior margin N. longicornis Thomson, 1879
7 Ommatidial part covering four-fifths of eye-stalk N. falklandensis Dahl, 1990
Ommatidial part not covering four-fifths of eye-stalk 8
8 Ommatidial part covering half of eye-stalk N. capensis Barnard, 1914
Ommatidial part covering two-thirds of eye-stalk 9
9 Denticles of pleonites 6 and 7 distally acute N. antarctica Dahl, 1990
Denticles of pleonites 6 and 7 rounded N. patagonica Dahl, 1990
10 Anterior margin of eye-stalk with 2 lobes 11
Anterior margin of eye-stalk with 3 lobes 12
11 Eye-stalk with flat anterior margin between lobes N. daytoni Vetter, 1996
Eye-stalk no flat margin between lobes N. schizophthalma Haney et al., 2001
12 Denticles of pleonites 6 and 7 acute distally N. troncosoi Moreira et al., 2003
Denticles of pleonites 6 and 7 rounded N. pseudotroncosoi Song et al., 2013
13 Shape of the eye-stalk sub-rectangular N. abyssicola Ledoyer, 1997
Shape of the eye-stalk oval 14
14 Ommatidial part covering half of the eye-stalk 15
Ommatidial part not covering half of the eye-stalk 20
15 Supraocular plate covering proximal portion of eye-stalk 16
Supraocular plate covering the half of eye-stalk N. deborahae Bochert & Zettler, 2012
16 Exopod of second maxilla clearly extend beyond the endopod of article 1 N. clausi Dahl, 1985
Exopod of second maxilla subequal length to endopod of article 1 17
17 Furcal rami almost the same length as combined length of telson and pleonites 6 and 7 N. marerubri Wägele, 1983
Furcal rami shorter than combined length of telson and pleonites 6 and 7 18
18 Furcal rami almost the same length as combined length of telson and pleonite 7 N. gerkenae Haney & Martin, 2000
Furcal rami longer than combined length of telson and pleonite 7 19
19 Article 4 of antennule with single distal spine N. brucei Olesen, 1999
Article 4 of antennule with one or two distal spines N. dahli Kazmi & Tirmizi, 1989
20 Ommatidial part covering more than two-thirds of eye-stalk 21
Ommatidial part covering more than three quarters of eye-stalk 32
21 Article 1 of endopod of second maxilla shorter than article 2 22
Article 1 of endopod of second maxilla not shorter than article 2 23
22 Rostrum length ca. 2.4 times as long as width N. lagartensis Escobar-Briones & Villalobos-Hiriart, 1995
Rostrum length ca. 1.9 times as long as width N. terazakii Othman et al., 2016
23 Article 1 of endopod of second maxilla subequal in length to article 2 N. kocatasi Moreira et al., 2007
Article 1 of endopod of second maxilla longer than article 2 24
24 Denticles of pleonites 6 and 7 square N. mortoni Lee & Bamber, 2011
Denticles of pleonites 6 and 7 not square 25
25 Denticles of pleonites 6 and 7 acute 26
Denticles of pleonites 6 and 7 rounded 27
26 Exopod of pleopod 1 bearing 15 or 16 spines on lateral margin N. melanophthalma Ledoyer, 2000
Exopod of pleopod 1 bearing 26 spines on lateral margin N. strausi Risso, 1826
27 Anal plates with distinct ‘shoulder’ N. bipes Dahl, 1985
Anal plates with no distinct ‘shoulder’ 28
28 Protopod of pleopod 4 with serrations along posterior margin 29
Protopod of pleopod 4 lacking serration along posterior margin 30
29 Article 3 of antennal peduncle bearing plumose setae on external lateral face N. mediterranea Kocak & Moreira, 2015
Article 3 of antennal peduncle lacking plumose setae on external lateral face N. kensleyi Haney & Martin, 2005
30 Furcal rami shorter than combined length of telson and pleonite 7 N. koreana Song et al., 2012
Furcal rami not shorter than combined length of telson and pleonite 7 31
31 Denticle of pleonite 2 with wide flat margin between denticles N. tagiri sp. nov., This paper
Denticle of pleonite 2 lacking flat margin between denticles N. reboredae Moreira et al., 2009
32 Ommatidial part covering four-fifths of eye-stalk N. villalobosi Ortiz et al., 2011
Ommatidial part covering three-quarters of eye-stalk 33
33 Denticles of pleonites 6 and 7 square-shaped N. ilheoensis Kensleyi, 1976
Denticles of pleonites 6 and 7 not squared 34
34 Denticles of pleonite 6 rounded and of pleonite 7 acute N. borealis Dahl, 1985
Denticles of pleonites 6 and 7 similar in shape 35
35 Denticles of pleonites 6 and 7 distally acute 36
Denticles of pleonites 6 and 7 rounded 37
36 Total length of mature female more than 10 mm, Exopod of pleopod 1 bearing more than 30 spines on lateral margin N. hessleri Martin et al., 1996
Total length of mature female less than 5 mm, Exopod of pleopod 1 bearing 15 or 16 spines on lateral margin N. neocaledoniensis Ledoyer, 2000
37 Article 2 of mandibular palp with single thin seta, anal plate with no distinct “shoulder” N. herbstii Leach, 1814
Article 2 of mandibular palp with two thin setae, anal plate with distinct “shoulder” N. dolsandoensis Song & Min, 2016

Acknowledgments

This study was supported by the Japan Agency for Marine-Earth Science and Technology (JAMSTEC). We are grateful to Dr. Lindsay D.J. of JAMSTEC, Dr. Victor S. Kuwahara of Soka University for reviewing the manuscript and making useful suggestions. We also thank the captain and crew of the R/V “Natsushima” and the operation team of the ROV “Hyper-Dolphin” for organizing the cruise and conducting the diving research and sampling, respectively. I am grateful to referees for their comments that helped improve the manuscript.

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