Research Article |
Corresponding author: Sook Shin ( shins@syu.ac.kr ) Academic editor: Alexander Martynov
© 2020 Michael Dadole Ubagan, Taekjun Lee, Philjae Kim, Sook Shin.
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:
Ubagan MD, Lee T, Kim P, Shin S (2020) A new species of the genus Henricia (Asteroidea, Spinulosida, Echinasteridae) from South Korea. ZooKeys 997: 1-15. https://doi.org/10.3897/zookeys.997.52703
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A new species of the genus Henricia Gray, 1840 that belongs to the family Echinasteridae is described from South Korea. Henricia epiphysialis sp. nov. has epiphyseal ossicles at the ends of the abactinal and lateral plates, and the abactinal and lateral spines form a hooked crown. The partial sequence of the mitochondrial COXI gene (537 bp) of H. epiphysialis sp. nov. was obtained, and the new species was morphologically and genetically compared with other related Henricia species.
Distribution, DNA barcoding, Henricia epiphysialis sp. nov., morphology, taxonomy
Echinasteridae Verrill, 1867 is the only family that belongs to the order Spinulosida Perrier, 1884. This family comprises eight accepted genera: Aleutihenricia Clark & Jewett, 2010; Dictyaster Wood-Mason & Alcock, 1891; Echinaster Müller & Troschel, 1840; Henricia Gray, 1840; Metrodira Gray, 1840; Odontohenricia Rowe & Albertson, 1988; Plectaster Sladen, 1889; and Rhopiella Fisher, 1940 (
A total of 11 species that belong to Aleutihenricia or Henricia have been reported in South Korea: Aleutihenricia beringiana Djakonov, 1950 and 10 Henricia species, namely, Henricia anomala Hayashi, 1973; Henricia elachys Clark & Jewett, 2010; Henricia leviuscula Stimpson, 1857; Henricia nipponica Uchida, 1928; Henricia ohshimai Hayashi, 1935; Henricia pachyderma Hayashi, 1940; Henricia pacifica Hayashi, 1940; Henricia regularis Hayashi, 1940; Henricia reniossa Hayashi, 1940; and Henricia sanguinolenta O.F. Müller, 1776 (
In DNA barcoding, sequence variation in a 658 bp region of the mitochondrial cytochrome c oxidase subunit I (COXI) gene is used for specimen identification and species discovery (
In this study, we identified a new species that belongs to the genus Henricia collected from waters adjacent to the East Sea, South Korea, and performed detailed morphological and molecular mitochondrial sequence analyses. This paper aims to extend the taxonomical insights to Henricia species in South Korea by providing a complete description of this new species.
In May and December 2014, sea stars were collected from the East Sea in South Korea by using fishing nets (Fig.
Total genomic DNA was isolated from ethanol-preserved tube feet tissue by using a DNeasy blood and tissue DNA isolation kit (Qiagen), according to the manufacturer instructions. The genomic DNA quality and concentration were determined using a Nanodrop ND-1000 spectrophotometer (Thermo Fisher Scientific). All genomic DNA samples were stored at −20 °C until further use. The partial sequence of the mitochondrial COI gene (658 bp) was amplified using a pair of primers, LCOech1aF1 (
List of Henricia species and GenBank accession numbers of COX1 gene used in this study.
Species | GenBank No. | Dataset | References | |
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189 bp | 537 bp | |||
H. compacta (Sladen, 1889) | KT268147 | + |
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H. epiphysialis sp. nov. | MT086587 | + | + | present study |
H. hayashii Djakonov, 1961 | LC336732 | + | + |
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H. hedingi Madsen, 1987 | KY853274 | + |
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H. kinkasana Hayashi, 1940 | LC336731 | + | + |
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H. leviuscula (Stimpson, 1857) | MK947912 | + | + | Lee and Shin 2019 |
H. lisa Clark, 1949 | KY853275 | + | + |
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H. nipponica Uchida, 1928 | LC336733 | + | + |
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H. obesa (Sladen, 1889) | KT268148 | + |
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H. oculata (Pennant, 1777) | KT268151 | + | + |
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H. ohshimai Hayashi, 1935 | LC336735 | + | + |
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H. ohshimai Hayashi, 1935 | LC336736 | + | + |
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H. pachyderma Hayashi, 1940 | MT079801 | + | + | Lee and Shin 2020 |
H. perforata (Müller, 1776) | KY853302 | + | + |
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H. pertusa (Müller, 1776) | KY853286 | + | + |
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H. regularis Hayashi, 1940 | LC336739 | + | + |
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H. reniossa Hayashi, 1940 | LC336740 | + | + |
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H. reticulata Hayashi, 1940 | LC336737 | + | + |
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H. sanguinolenta (Müller, 1776) | HM542200 | + |
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H. sanguinolenta (Müller, 1776) | KY853253 | + | + |
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H. spongiosa (Fabricius, 1780) | KY853268 | + | + |
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H. tumida Verrill, 1909 | LC336747 | + | + |
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Henricia sp. 1 | LC336744 | + | + |
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Henricia sp. 2 | LC336742 | + | + |
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Henricia sp. 3 | LC336743 | + | + |
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Henricia sp. 4 | LC336741 | + | + |
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Henricia sp. 5 | LC336738 | + | + |
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Henricia sp. 6 | LC336745 | + | + |
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Henricia sp. 7 | LC336746 | + | + |
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Henricia sp. 8 | LC336730 | + | + |
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Henricia sp. 9 | KY853310 | + | + |
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Echinaster brasiliensis Müller & Troschel, 1842 | MG636999 | + | + | Seixas et al. 2018 |
E. callosus Marenzeller, 1895 | KT268121 | + |
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E. luzonicus (Gray, 1840) | KT268137 | + |
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E. sepositus (Retzius, 1783) | LC336729 | + | + |
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Class Asteroidea de Blainville, 1830
Order Spinulosida Perrier, 1884
Family Echinasteridae Verrill, 1867
Genus Henricia Gray, 1840
Holotype : South Korea • 1 specimen; waters adjacent to Namae, 37°55'57.31"N, 128°48'45.58"E; 40 m; 28 May 2014; S. Shin and T. Lee; fishing net; MERBK–A–1255. Paratypes: South Korea • 1 specimen; waters adjacent to Jukbyeon, 37°3'32.49"N, 129°26'14.57"E; 100 m; 19 Dec. 2014; S. Shin and T. Lee; fishing net; NIBRIV0000837785. 1 specimen; waters adjacent to Namae, 37°55'57.31"N, 128°48'45.58"E; 40 m; 28 May 2014; S. Shin and T. Lee; fishing net; MERBK–A–1256.
Regular size, R/r = 4.9–5.4, abactinal plates crowded with 11–40 spines, abactinal and lateral spines forming distinct hooked crown, epiphyseal ossicles formed at ends of abactinal and lateral plates, one to three papulae, marginal and ventrolateral series distinguishable, adambulacral plates bearing 10–14 slender spines.
Holotype. (Figs
Arms five, semi-cylindrical, gradually tapering to tips (Fig.
External characteristics of Henricia epiphysialis sp. nov. A abactinal side B actinal side C abactinal spines D spines on lateral side of arm E adambulacral spines F oral part. Abbreviations: ab abactinal side ls lateral side ss superomarginal spines is inferomarginal spines vs ventrolateral spines as adambulacral spines os oral spines ms marginal spines sos sub-oral spines.
Paratypes. Size. R = 39 mm, r = 8 mm, R/r = 4.9; R = 60 mm, r = 11 mm, R/r = 5.4.
The specific name is derived from the Latin “epiphysialis,” which means the end part of a long bone.
Denuded skeleton of Henricia epiphysialis sp. nov. A abactinal plates B madreporite (arrow) C part of abactinal and lateral side of arm D plates on the lateral and actinal side of arm E actinal plates F oral part. Abbreviations: ab abactinal side ls lateral side eo epiphyseal ossicles p papula s superomarginal plates in intermarginal plates i inferomarginal plates v ventrolateral plates a adambulacral plates am ambulacral plates o oral plates.
This species is found on hard substrates (rocky bottom) from a shallow water of a depth of 40 m to 100 m.
We determined the phylogenetic relationships based on two COX1 datasets with 27 and 31 species of the genus Henricia respectively, including H. epiphysialis sp. nov., on the basis of 189 and 537 bp of the mitochondrial COX1 gene by using the NJ method. All Henricia species formed a monophyletic group with congeneric species and were clearly distinguished from the sister taxa (Fig.
Comparison of the morphological characteristics of Henricia epiphysialis sp. nov. with related Henricia species. Morphological data derived from the respective original descriptions, the present study, and
Henricia epiphysialis sp. nov. | H. compacta (Sladen, 1889) | H. densispina (Sladen, 1878) | H. djakonovi Chichvarkhin, 2017 | H. exigua Hayashi, 1940 | H. kinkasana Hayashi, 1940 | H. leviuscula (Stimpson, 1857) | H. regularis Hayashi, 1940 | H. reniossa Hayashi, 1940 | H. skorikovi Djakonov, 1950 | |
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R/r | 4.9–5.4 | 4.4–7.5 | 2.9–5.5 | 4.7–5.0 | 4.2–4.5 | 4.0–4.8 | 5.0–6.0 | 3.1–5.0 | 5.5–8.0 | 3.6–5.0 |
Number of spines of abactinal plates | 11–40 | up to 45 | 30 or more | 20–30 | 5–13 | 5–18 | 40–60 | 9–20 | 15–60 or more | up to 16 |
Shape of abactinal spines | hooked crown | stout | granular | stout, barrel, bullet tip | slender, pointed tip |
fine, slender, pointed tip | short, granular, solid glassy tip |
slender, pointed tip |
very fine, rough tip |
short, robust, thorny tip |
Number of abactinal papulae | 1–3 | 1–3 | 1–3, rarely 5 | 1 or 2 | 1–3 | single | 1–3 | single | 1 or 2 | 2–6 |
Shape of abactinal plates | reniform with conspicuous epiphyseal ossicle |
crescentic, compact |
subtriangular | cross, oval, triangular, irregular | elliptic, quasi-triangular, quasi-quadrate |
crescentic | elliptic, reniform, subquadrate | subquadrate | reniform | slender, rod-like |
Shape of actinal plates | rounded cross, elongated cross | quadrilobed, squarish | elongated cross | square pillow | rounded cross, elongated cross | rounded cross, elongated cross | elongated cross, small rod-like | rounded cross | elongated cross, quasi-quadrate | elongated cross, rod-like |
Number of actinal papulae | single | single | single | single | single | single | single | single | 1 or 2 | unknown |
Number of adambulacral spines | 10–14 | 5 or 6 | 11–16 | 10 or 11 | 13–15 | 8–12 | 15–18 | 9–13 | 15–25 | 7–12 |
Number of furrow spines | single | 2 or 3 | single | single | double | single | single | single or double | single | single |
Pattern of adambulacral spines (near ambulacral furrow + near ventrolateral plate) | 1 long, 2 slender, bluntly pointed + 4–14 slightly shorter |
2 or 3 prominent + 4–6 slightly shorter | 2 or 3 prominent, bluntly pointed + 4–16 slightly shorter | 2 or 3 larger + 4–11 slightly shorter | 1 long, 2 spatulate + 4–15 shorter | 1 long, 2 slender, bluntly pointed + 4–12 slightly shorter | 1 long, 3 stout + 5–18 slightly shorter |
1 long, 2 slender, bluntly pointed + 4–13 slightly shorter | 1 long, 5 slender + 7–25 slightly shorter |
1 long, 2 coarse + 4–12 slightly shorter |
Distribution | Korea (East Sea) | southern Australia | Bohai Sea, Yellow Sea, Korea Strait, Tatar Strait, Kurile Island, Japan, Philippines | Rudnaya Bay, Kievka Bay | southern Japan, East China Sea | Japan (off Kinkasan) | Korea (East Sea), Alaska (Kadiak) | East China Sea, Korea (East Sea, Korea Strait, Jeju Island), Japan (Goto Island, Uraga Channel) | Korea (East Sea), Japan (Yezo Strait) | White Sea, Barents Sea, Chesha Bay |
Pairwise genetic comparison for 537 bp of the mitochondrial COX1 gene in 27 species of Henricia including Henricia epiphysialis sp. nov.
Species | 1 | 2 | 3 | 4 | 5 | 6 | 7 | 8 | 9 | 10 | 11 | 12 | 13 | 14 | 15 | 16 | 17 | 18 | 19 | 20 | 21 | 22 | 23 | 24 | 25 | 26 | 27 | 28 |
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H. epiphysialis sp. nov. | ||||||||||||||||||||||||||||
H. hayashii | 0.069 | |||||||||||||||||||||||||||
H. kinkasana | 0.065 | 0.056 | ||||||||||||||||||||||||||
H. leviuscula | 0.141 | 0.143 | 0.132 | |||||||||||||||||||||||||
H. lisa | 0.084 | 0.069 | 0.054 | 0.114 | ||||||||||||||||||||||||
H. nipponica | 0.096 | 0.114 | 0.112 | 0.157 | 0.096 | |||||||||||||||||||||||
H. oculata | 0.131 | 0.143 | 0.134 | 0.150 | 0.139 | 0.161 | ||||||||||||||||||||||
H. ohshimai 1 | 0.129 | 0.143 | 0.141 | 0.149 | 0.136 | 0.150 | 0.064 | |||||||||||||||||||||
H. ohshimai 2 | 0.142 | 0.166 | 0.151 | 0.161 | 0.159 | 0.170 | 0.079 | 0.094 | ||||||||||||||||||||
H. pachyderma | 0.154 | 0.161 | 0.159 | 0.160 | 0.154 | 0.188 | 0.136 | 0.145 | 0.133 | |||||||||||||||||||
H. perforata | 0.152 | 0.164 | 0.152 | 0.154 | 0.167 | 0.165 | 0.117 | 0.141 | 0.138 | 0.143 | ||||||||||||||||||
H. pertusa | 0.067 | 0.048 | 0.036 | 0.136 | 0.061 | 0.105 | 0.141 | 0.138 | 0.151 | 0.173 | 0.166 | |||||||||||||||||
H. regularis | 0.090 | 0.083 | 0.071 | 0.145 | 0.085 | 0.118 | 0.152 | 0.138 | 0.165 | 0.188 | 0.191 | 0.065 | ||||||||||||||||
H. reniossa | 0.077 | 0.058 | 0.056 | 0.114 | 0.042 | 0.096 | 0.136 | 0.134 | 0.159 | 0.154 | 0.169 | 0.052 | 0.083 | |||||||||||||||
H. reticulata | 0.135 | 0.133 | 0.133 | 0.171 | 0.152 | 0.173 | 0.120 | 0.122 | 0.113 | 0.131 | 0.159 | 0.133 | 0.146 | 0.147 | ||||||||||||||
H. sanguinolenta | 0.084 | 0.067 | 0.056 | 0.118 | 0.013 | 0.092 | 0.139 | 0.134 | 0.159 | 0.154 | 0.164 | 0.063 | 0.090 | 0.034 | 0.150 | |||||||||||||
H. spongiosa | 0.088 | 0.071 | 0.061 | 0.118 | 0.017 | 0.092 | 0.144 | 0.139 | 0.164 | 0.159 | 0.169 | 0.067 | 0.094 | 0.038 | 0.154 | 0.004 | ||||||||||||
H. tumida | 0.104 | 0.117 | 0.115 | 0.161 | 0.124 | 0.135 | 0.137 | 0.137 | 0.146 | 0.156 | 0.165 | 0.120 | 0.133 | 0.124 | 0.128 | 0.124 | 0.129 | |||||||||||
Henricia sp. 1 | 0.090 | 0.087 | 0.077 | 0.141 | 0.071 | 0.086 | 0.150 | 0.145 | 0.162 | 0.150 | 0.156 | 0.081 | 0.111 | 0.058 | 0.160 | 0.067 | 0.071 | 0.133 | ||||||||||
Henricia sp. 2 | 0.062 | 0.062 | 0.044 | 0.120 | 0.059 | 0.105 | 0.141 | 0.145 | 0.158 | 0.159 | 0.154 | 0.050 | 0.081 | 0.056 | 0.147 | 0.056 | 0.061 | 0.113 | 0.077 | |||||||||
Henricia sp. 3 | 0.058 | 0.052 | 0.015 | 0.123 | 0.052 | 0.105 | 0.136 | 0.136 | 0.156 | 0.156 | 0.159 | 0.032 | 0.069 | 0.048 | 0.140 | 0.055 | 0.059 | 0.122 | 0.073 | 0.040 | ||||||||
Henricia sp. 4 | 0.063 | 0.054 | 0.006 | 0.129 | 0.052 | 0.110 | 0.131 | 0.138 | 0.149 | 0.156 | 0.154 | 0.034 | 0.069 | 0.054 | 0.137 | 0.055 | 0.059 | 0.113 | 0.075 | 0.042 | 0.009 | |||||||
Henricia sp. 5 | 0.058 | 0.048 | 0.015 | 0.127 | 0.050 | 0.103 | 0.141 | 0.138 | 0.154 | 0.152 | 0.164 | 0.029 | 0.058 | 0.048 | 0.138 | 0.053 | 0.057 | 0.120 | 0.073 | 0.044 | 0.011 | 0.013 | ||||||
Henricia sp. 6 | 0.056 | 0.048 | 0.015 | 0.123 | 0.046 | 0.099 | 0.136 | 0.134 | 0.149 | 0.149 | 0.164 | 0.029 | 0.058 | 0.044 | 0.140 | 0.048 | 0.053 | 0.117 | 0.069 | 0.040 | 0.011 | 0.013 | 0.004 | |||||
Henricia sp. 7 | 0.058 | 0.050 | 0.017 | 0.125 | 0.048 | 0.096 | 0.134 | 0.131 | 0.147 | 0.147 | 0.161 | 0.031 | 0.061 | 0.046 | 0.138 | 0.050 | 0.055 | 0.120 | 0.067 | 0.042 | 0.013 | 0.015 | 0.006 | 0.002 | ||||
Henricia sp. 8 | 0.141 | 0.145 | 0.136 | 0.131 | 0.141 | 0.147 | 0.096 | 0.107 | 0.111 | 0.145 | 0.107 | 0.138 | 0.147 | 0.138 | 0.143 | 0.134 | 0.138 | 0.147 | 0.138 | 0.141 | 0.141 | 0.134 | 0.136 | 0.132 | 0.129 | |||
Henricia sp. 9 | 0.145 | 0.143 | 0.145 | 0.148 | 0.157 | 0.176 | 0.107 | 0.127 | 0.139 | 0.147 | 0.129 | 0.154 | 0.166 | 0.150 | 0.105 | 0.152 | 0.157 | 0.146 | 0.168 | 0.145 | 0.138 | 0.138 | 0.148 | 0.148 | 0.150 | 0.125 | ||
E. brasiliensis | 0.220 | 0.212 | 0.228 | 0.210 | 0.217 | 0.207 | 0.227 | 0.217 | 0.202 | 0.190 | 0.200 | 0.230 | 0.230 | 0.233 | 0.202 | 0.220 | 0.225 | 0.197 | 0.217 | 0.227 | 0.225 | 0.230 | 0.225 | 0.223 | 0.220 | 0.205 | 0.217 | |
E. sepositus | 0.192 | 0.190 | 0.207 | 0.190 | 0.207 | 0.190 | 0.192 | 0.195 | 0.183 | 0.214 | 0.192 | 0.207 | 0.215 | 0.210 | 0.219 | 0.205 | 0.205 | 0.204 | 0.209 | 0.190 | 0.210 | 0.205 | 0.210 | 0.205 | 0.205 | 0.178 | 0.212 | 0.190 |
The range and average p-distance values of Henricia species examined in this study.
Group | Range | Average |
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H. epiphysialis–other Henricia sp. | 0.056–0.154 | 0.098 |
All of Henricia species | 0.002–0.188 | 0.110 |
All Henricia species except for H. kinkasana and Henricia sp.3–sp.7 | 0.040–0.188 | 0.114 |
Phylogenetic trees of Henricia species including Henricia epiphysialis sp. nov. based on Neighbor joining (NJ) A NJ tree constructed with 189 bp of COX1 dataset B NJ tree constructed with 537 bp of COX1 dataset; K,
The diagnostic combination of the morphological characteristics of the genus Henricia such as spination of the abactinal (primarily on the disk and proximal portion of the arm), adambulacral, and oral plates was found to be consistent and reliable for determining the species (e.g.,
Other morphological characteristics of H. epiphysialis sp. nov., such as the ratio of arm to disk and number of adambulacral spines, are similar to those of H. kinkasana which is a slender-rayed species; however, this new species differs mainly in the number of abactinal spines and shape of both abactinal and lateral spines. Henricia epiphysialis sp. nov. has 11–40 robust abactinal spines on the abactinal plate, whereas H. kinkasana has five to 18 fine, delicate abactinal spines. Moreover, the conspicuous epiphyseal ossicles at the ends of the abactinal and lateral plates are exclusively present in H. epiphysialis sp. nov. Therefore, the extension of ossicles in the plate and hooked crown shape of the spines are diagnostic characteristics for this new species.
In this study, we identify a new Henricia species based on its morphological characteristics and DNA barcoding. Henricia epiphysialis sp. nov. has distinct morphological features and was classified as a new species after comparison with related species. Moreover, the molecular analysis showed that H. epiphysialis sp. nov. clearly formed a monophyletic node in a large clade of the genus Henricia species (Fig.
This study was supported by the project titled “Improvement of management strategies on marine disturbing and harmful organisms” funded by the Ministry of Oceans and Fisheries, South Korea (No. 20190518) and the Marine Biotechnology Program of the South Korea Institute of Marine Science and Technology Promotion (KIMST) funded by the Ministry of Oceans and Fisheries (MOF) (No. 20170431), and a grant from the National Institute of Biological Resources (NIBR), which was funded by the Ministry of Environment of the Republic of South Korea (NIBR202002110).