Research Article |
Corresponding author: Takuma Hirata ( fullkamuhira@gmail.com ) Academic editor: Tammy Horton
© 2019 Takuma Hirata, Yoshihiro Fujiwara, Tomohiko Kikuchi.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Hirata T, Fujiwara Y, Kikuchi T (2019) A new species of Nebalia (Crustacea, Leptostraca) from a hydrothermal field in Kagoshima Bay, Japan. ZooKeys 897: 1-18. https://doi.org/10.3897/zookeys.897.37061
|
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.
Hydrothermal vent, key, Leptostraca, Malacostraca, Nebalia, new species
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
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.
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.
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.
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).
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).
Female holotype. Carapace (Fig.
Rostrum
(Fig.
Compound eye
(Fig.
Antennule
(Fig.
Antenna
(Fig.
(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.
First maxilla
(Fig.
Second maxilla
(Fig.
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.
Thoracopod 2–6
(Fig.
Thoracopod 7
(Fig.
Thoracopod 8
(Fig.
Pleon
(Fig.
Pleopod 1
(Fig.
Pleopod 2
(Fig.
Pleopod 3
(Fig.
Pleopod 4
(Fig.
Pleopod 5
(Fig.
Pleopod 6
(Fig.
Telson, anal plates, and furca
(Fig.
Color in life
(Fig.
(adult male): antennule flagellum more swollen proximally than in female (Fig.
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.
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.
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.
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 |
|
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 |
|
N. borealis | Sand | 240 m | 2.1 times | Oval | 2 sp | – | – | – | 1 ts | 24 sp | ≥ Pn7+t | Round to acute | Distinct |
|
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 |
|
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 |
|
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 |
|
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
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.
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 |
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.