The First Larval Description of the Common Aesop Slipper Lobster Scyllarides haanii (Decapoda: Achelata: Scyllaridae)

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Species Diversity 29: 327–335 The First Larval Description of the Common Aesop Slipper Lobster Scyllarides haanii (Decapoda: Achelata: Scyllaridae) Kooichi Konishi1,3, Takashi Yanagimoto1, and Seinen Chow2 1 Fisheries Resources Institute, Japan Fisheries Research and Education Agency, Yokohama, Kanagawa 236-8648, Japan E-mail: [email protected] (KK) 2 Aquos Institute, 3-2153-79 Motohachioji, Hachioji, Tokyo 193-0826, Japan 3 Corresponding author (Received 19 March 2024; Accepted 26 August 2024) A late-stage phyllosoma, collected from the North Pacific Ocean, was identified as Scyllarides haanii (De Haan, 1841) by DNA analysis. A morphological description of the phyllosoma of this common Indo-Pacific species is given for the first time. Larval characteristics of late phyllosoma stages are compared with those of previously described Scyllarides Gill, 1898 and related genera. A tentative key to scyllarid phyllosomas in the late-stages is also provided. Key Words: Crustacea, Achelata, Scyllaridae, phyllosoma larva, DNA barcoding. Introduction The genus Scyllarides Gill, 1898 in the subfamily Arctidinae consists of 14 species (Holthuis 1991; DecaNet 2023), and larval descriptions have been given in 7 species (Table 1). It is notable that the phyllosoma stages of S. haanii (De Haan, 1841) are yet to be described in the remaining 9 species of Scyllarides, despite the wide distribution of adult lobsters in the Pacific and Indian Oceans (Morin and MacDonald 1984; Holthuis 1991; Báez et al. 2022). A latestage scyllarid phyllosoma specimen, collected from the western North Pacific Ocean, was examined morphologically and identified as S. haanii by DNA barcoding. We present a morphological description of the larval specimen and compare the larval characteristics of Scyllarides species with those of related genera. Materials and Methods In the research cruise for spawning stock of neon squid Ommastrephes bartramii (Lesueur, 1821) in the North Pacific Ocean performed by R/V Kaiyo-Maru, Fisheries Agency of Japan, a large scyllarid phyllosoma was caught using a ring net (2 m mouth opening, 526 µm mesh) on 30 October 2013 at 33°01′N, 156°59′E. The net was towed on the surface at a speed of 1.9 kt during daytime (10:30–10:56 hours). This phyllosoma was fixed in 80% ethanol on board and transferred to the Fisheries Resources Institute. A small tissue piece was dissected from a pereiopod, washed well with tap water, and homogenized, from which crude DNA was extracted using a DNA extraction kit (QuickGene DNA tissue kit, DT-S, KURABO). Amplification protocols of partial mitochondrial 16S rDNA sequence and nucleotide sequence determination were followed by Ueda et al. (2021). The nucleotide sequence determined was subjected to BLAST (basic local alignment search tool) (Altshul et al. 1990) in National Center for Biotechnology Information to find identical or similar sequences. Nucleotide sequence alignment was performed using the ClustalW algorithm implemented in MEGA 6 (Tamura et al. 2013) followed by manual editing. Calculation of K2P distance between sequences, selecting the best-fit model of nucleotide substitution, and construction of a neighbor-joining phylogenetic tree was performed using MEGA 6. After the molecular analyses, the appendages were dissected using fine insect pins. Observations and drawings were made with the aid of a drawing tube attached to an Olympus BX51 microscope and an SZX10 stereomicroscope. The measurements were taken as follows: total length (TL) from the anterior margin of the cephalic shield between the eyes to the posterior margin of the telson, cephalic shield length (CL) from the anterior to the posterior margin of the cephalic shield, pleon length (PL) from the base of pleon to the posterior telsonal margin, cephalic shield width (CW) and thorax width (TW) measured at the widest part of the cephalic shield and thorax, respectively. The voucher specimen of the phyllosoma, KY13-5-102 in the serial number by Fisheries Agency research cruise, was deposited at the Hokkaido University Museum under the catalog number ICHUM-8558. Results DNA analysis. The nucleotide sequence determined was 432 bp and available in the public databases through the DNA Data Bank of Japan under the International Nucleotide Sequence Databases (INSD) accession numPublished online 10 October 2024 DOI: 10.12782/specdiv.29.327 328 Kooichi Konishi et al. ber LC763119. Against this sequence, the most similar sequences returned by BLAST search were of Scyllarides haanii (INSD accession numbers JN701690, JN701691, and MN817127), in which the K2P distances between the sequences ranged 0.00%–1.17%. The phylogenetic tree (Fig. 1) unambiguously determined this phyllosoma to be S. haanii. Larval description of Scyllarides haanii, stage X. The phyllosoma stage of the present specimen was determined according to Robertson’s (1969a) stage key for phyllosoma. Dimensions: TL = 39.5 mm, CL = 28.8 mm, CW = 25.0 mm, TW = 17.8 mm, PL = 6.4 mm. Cephalothorax (Figs 2, 3A): outline of cephalic shield ellipsoid, slightly wider than long (CL/CW ratio = 1.15), and much wider than thorax (CW/TW ratio = 1.40). Small overhang plate behind the base of eyestalk (Fig. 3B). Antennule (Fig. 4A): peduncle 3-segmented, outer flagellum longer than half length of inner flagellum, with 5–6 tiers of aesthetascs on its inner margin. Antenna (Fig. 4B): flattened dorsoventrally, 3-segmented, the second segment anteriorly pointed. Mandibles (Fig. 4C): slightly flattened dorsoventrally, asymmetrical in dentition. Incisor process and medial gnathal edge with a series of teeth. Molar process crowned by many denticules and minute papillae. Distal inner half portion of mandible covered with labrum and paragnath. Maxillule (Fig. 4D): no endopod projection. Basal endite with 3 stout spines and 8 subterminal setae. Coxal endite with 12–13 stout setae. Maxilla (Fig. 4E): endopodal area with 2 thin setae. Basal and coxal area with 2 and 1 thin anterior setae, respectively. Scaphognathite without marginal plumose setae. Maxilliped 1 (Fig. 4F): 3-lobed process, with a short seta on endopodal area. Maxilliped 2 (Fig. 4G): endopod 5-segmented, penultimate segment with 2 stout spines and 12 setae in total. Exopod rudimentary bud. No gill rudiments. Maxilliped 3 (Fig. 4H): endopod 4-segmented, with many setae on distal segment. Exopod rudimentary bud. No gill rudiments. Pereiopods 1–5 (Fig. 3A): exopods distally flagellated with annuli bearing a pair of natatory setae. No gill rudiments on pereiopods and thorax pleura. Pleon (Fig. 3C): much narrower than thorax, somite segmented with biramous rudiments of pleopods (Fig. 3D) on somites 2–5. Uropod biramous, posterior margins of endopod and exopod reaching that of the telson. Telson oval without spines. Discussion Of the 17 species in the subfamily Arctidinae includTable 1. Previous larval descriptions in the genus Scyllarides. Species Source of materials Collection locality Described stages References S. aequinoctialis L/P Florida, Bahama, Columbia I–XI 1 S. aequinoctialisaP Gulf of Mexico I–VI 2a S. aequinoctialis P Western Atlantic IV 3 S. astoribP Gulf of California I, III, IV, VII 4 S. astori P Gulf of California Late phyllosoma stages 5 S. astori L Gulf of California I 6 S. astori P Gulf of California ni 7 S. elisabethaecP Off Durban Late phyllosoma stages 8 S. herklotsii P St. Helena Island ni 9 S. herklotsii P/L Gulf of Guinea ns, I, III, V, VI, VIII, IX, ni 10 S. latus L Mediterranean I 11 S. latus P Mediterranean II 12 S. latus P Mediterranean I 13 S. nodifer PAtlantic Late phyllosoma stages 1 S. squammosus P New Caledonia Late phyllosoma stages, ni 14 S. squammosus L Off Kagoshima I–VId15 S. squammosus P Western Pacific Late phyllosoma stages 16 S. squammosus L New Caledonia I 17 S. squammosus P*Coral Sea VI–XII 18 S. squammosus P*Off Ogasawara Islands ni 19 S. squammosus P*Northwest Pacific IX–XI 20 S. sp. P*Northwest Pacific VII, IX 20 S. sp-B P*Central North Pacific IX–XI 20 S. sp-C P*eNorthwest Pacific XII 20 Abbreviations: L, laboratory-reared; ni, nisto; ns, naupliosoma; P, plankton collection or wild-caught; P*, identified by DNA analyses. a In their paper, the figure captions 4 and 5 are placed reversely; b as ‘phyllosoma X’; c as Species Z; d instar number; e DNA was analyzed but species could not be determined. References: 1 Robertson (1969a); 2 Manzanilla-Dominguez and Gasca (2004); 3 Santana et al. (2007); 4 Johnson (1968); 5 Johnson (1970); 6 Johnson and Knight (1975); 7 Johnson (1975); 8 Berry (1974); 9 Chace (1966); 10 Crosnier (1972); 11 Santucci (1925); 12 Fiedler and Spanier (1996); 13 Torres et al. (2013); 14 Michel (1968); 15 Saisho and Sone (1971); 16 Sekiguchi (1990); 17 Coutures (2001); 18 Palero et al. (2016); 19 Hidaka et al. (2022); 20 Chow et al. (2022a). Larval description of Scyllarides haanii 329 ing Scyllarides, larval description has been given for 9 species in 2 genera, representing 53% of the total. However, in most of these morphological descriptions of larvae, their parentage was based on estimation between locations of planktonic specimens and presumed adult distributions. Sometimes this fact can be controversial for decapod larval studies, concerning plankton dynamics. For example, Chow et al. (2006) commented that long-lived teleplanic larvae such as phyllosoma may drift to distant waters due to the long planktonic period, where their adult forms do not exist. The accurate specific identification of phyllosomas by more certain methods, such as laboratory-rearing or DNA barcoding, is essential for basic population studies and stock management. To date, only four cases have been known for the accurate parentage determination in the arctidine species; Arctides regalis Holthuis, 1963, S. aequinoctialis (Lund, 1793), S. herklotsii (Herklots, 1851), and S. squammosus (H. Milne Edwards, 1837) (Robertson 1969a; Crosnier 1972; Palero et al. 2016; Chow et al. 2021). The present study is the fifth larval description based on the accurate parentage in the Arctidinae. According to Robertson’s (1969a) stage key for S. aequinoctialis, the present specimen is assigned to stage X in pleon extension beyond level of origin of pereiopod 5 and in lack of gill buds on pereiopods. However, in the body size, TL of S. haanii is about 1.4 times larger than S. aequinoctialis in the stage X. This size is near to Sims’ (1965) final XII stage, which was originally described as ‘Parribacus’ and later argued assignation to Scyllarides (Robertson 1969b; Sekiguchi 1988). In Robertson’s (1969a) phyllosoma specimens, the earlyto mid-stages were obtained by laboratoryrearing while the late-stages were collected from plankton. The accuracy of species identification in its lateto finalstages may be debatable. As only one specimen was available in our study, it is currently unclear whether the difference in body size between S. aequinoctialis and S. haanii is due to specific characteristics or not. In the preceding larval works, two groups have been recognized for the midto late-stage phyllosomas of the Scyllarides (Robertson 1969a; Sekiguchi 1986, 1988)— the first group: S. herklotsii, S. nodifer (Stimpson, 1866), S. squammosus; and the second group: S. aequinoctialis, S. astori (Holthuis, 1963). The morphology of the present study shows the phyllosoma of S. haanii belongs to the second group. In the Arctidinae, three morphotypes are categorized in the late-stage phyllosomas as follows: Type 1: cephalic shield narrow ovoid, and posterior margin of thorax concave between pereiopod 5 with setose exopod—Arctides Holthuis, 1960. Type 2: cephalic shield ellipsoid, and posterior margin of thorax concave between pereiopod 5 without setose exopod—S. herklotsii, S. nodifer, S. squammosus. Type 3: cephalic shield ellipsoid, and posterior margin of thorax concave between pereiopod 5 with setose exopod—S. aequinoctialis, S. astori, S. elisabethae (Ortmann, 1894), and possibly S. haanii. Type 2 and Type 3 also correspond to Sekiguchi’s (1988) first and second groups of Scyllarides phyllosoma, respectively. In the family Scyllaridae, the late-stage phyllosomas of the genus Parribacus Dana, 1852 in the subfamily Ibachinae show a close resemblance to those of Scyllarides (Phillips et al. 1981; Palero et al. 2014a; Chow et al. 2022b). In the Parribacus, however, the second antenna of midto late-stage phyllosomas are unsegmented, whereas those of the Arctidinae possess segmented antennae. Furthermore, the cephalic shield outline is slightly obovoid in Parribacus. Even in the Parribacus phyllosomas, antennal segmentation occurs in the final stage (or gill stage), suggesting that the discrepancy observed can be attributed to variations in the developmental rates among different taxonomic groups. Our present result also confirms a close resemblance of the late-stage Parribacus phyllosomas to those of Scyllarides. To summarize the preceding arguments, Table 2 provides a comparison of the main characteristics of phyllosoma at or near the same developmental stage with those previously described in the Arctidinae and Parribacus of the Ibacinae. In this instance, our objective is to make comparisons between larval characteristics at the same developmental stage wherever possible, given the significant morphological changes that occur during phyllosoma development. Baisre (1994) showed a close relationship between Arctides, Scyllarides and Parribacus, typified by the phyllosomas. Chow and Yanagimoto (2021) and Chow et al. (2022a) stated as a general feature of late-stage phyllosoma larvae Fig. 1. Neighbor-joining phylogenetic tree drawn based on Kimura-2-parameter distances of partial 16S sequences between 11 arctidine slipper lobster species and the sculptured slipper lobster [Parribacus antarcticus (Lund, 1793) and P. japonicus Holthuis, 1960] as outgroup species. Accession numbers are shown in parentheses. Bootstrap values of > 50% (out of 1000 replicates) are shown at the nodes. The same tree topology was obtained by maximum likelihood and maximum parsimony methods. 330 Kooichi Konishi et al. Fig. 2. Scyllarides haanii (De Haan, 1841), stage X phyllosoma (ICHUM-8558). Whole body in dorsal view. Scale bar = 10 mm. Fig. 3. Scyllarides haanii (De Haan, 1841), stage X phyllosoma (ICHUM-8558). A, Whole body in ventral view; B, anterior part of cephalon in dorsal view; C, pleon in dorsal view; D, right pleopod 5. Larval description of Scyllarides haanii 331 that the depression between the base of pereiopod 4 is posterior to the thorax in arctidine phyllosomas, whereas it is the base of pereiopod 5 in those of the ibachine genus Parribacus. However, their diagnostic larval characteristics are not a key to completely distinguishing these two genera, as there are two larval morphotypes in Scyllarides phyllosomas. Although Parribacus phyllosomas in the late stages resemble Type 3 listed above, these are distinguished by the antenna with no or fewer segments than those of Scyllarides. Additionally, the Parribacus phyllosomas reach the largest body size in the Achelata; TL of the final stage phyllosoma ranging 70–80 mm, which is larger than 1.6 times those of arctidine phyllosomas. The present result reconfirms Scyllarides phyllosomas consist of at least two morphotype groups in the posterior thoracic marginal depression between pereiopods and exopods of pereiopod 5. Phillips et al. (1981) suggested that their ‘Scyllarides sp. a’ was similar to the final stage phyllosoma of S. squammosus known at that time. Sekiguchi (1990) noted that ‘Scyllarides sp. a’ was distinguished from S. squammosus and possibly the final stage of S. haanii. The final stage of S. squammosus by Palero et al. (2016) also well corresponds with ‘Scyllarides sp. a’ in the antenna, the dactylus on pereiopods, and the mid-dorsal posterior spine on the pleonal somites 2–5. Our present result on S. haanii supports that the ‘Scyllarides sp. a’ is the final phyllosoma of S. squammosus. The dactylus length of the pereiopod, i.e., short hook-shaped or long sickle-shaped, may also be complementary characteristic. Fig. 4. Scyllarides haanii (De Haan, 1841), stage X phyllosoma (ICHUM-8558). A, Antennule; B, antenna; C, left mandible in ventral view; D, maxillule (left); E, maxilla (right); F, maxilliped 2; G, maxilliped 3 showing rudimentary exopod (arrow). 332 Kooichi Konishi et al. However, as is well known, this fragile part is almost lacking in the field-collected specimens. Three unidentified phyllosomas, ‘S. sp.’, ‘S. sp-B’, and ‘S. sp-C’ by Chow et al. (2022a) belong to Type 2 in this study. Johnson (1971) attributed a series of mid-late phyllosomas collected in the Hawaiian waters to S. squammosus. Although S. squammosus and S. haanii coexist in this area (Holthuis 1991), these phyllosomas are clearly distinguished by morphotype, i.e., Types 2 and 3, and Johnson’s (1971) specific identification is now confirmed. In addition, the giant scyllarid phyllosoma reported by Kishinouye (1918) from the stomach contents of the yellowfin tuna, 45 mm in body length, is the final stage of Scyllarides phyllosoma which belongs to Type 2. The unidentified giant scyllarid phyllosoma specimens described by Sims (1964) as ‘Phyllosoma A–D’ from the Florida Straits appears to belong to lateand final-stages of Scyllarides, based on the characteristics as follows: 1) cephalic shield slightly wider than the thorax (A–D), 2) posterior margin of thorax concave between pereiopod 5 (A–D), 3) pereiopods without subexopodal spines (A–D), 4) short dactylus on pereiopod 5 (B and C), and 5) pereiopod 5 with setose exopod (A, B, and D). In the adult classification of the Scyllaridae, close affinity among three Scyllarides species, S. haanii, S. latus, and S. squammosus was confirmed by DNA analyses (Liu and Wang 2020). Within these three species, our result shows that the phyllosoma morphology of S. haanii differs from the remainder at least in the late stages. Nevertheless, the reliability of the parentage within the Arctidinae remains significantly incomplete not only in larval morphology but also in the context of DNA phylogenetic analyses even in Scyllarides. The discrepancy between the results of larval morphotypes and those of adult groups based on genetic analyses and/or laboratory rearing should be addressed with further data. The CW/TW ratio may be useful for species discrimination from mid to late stages in some cases (Chow et al. 2022a, b), but this morphometric characteristic is not necessarily universal for all developmental stages because they can change from the late to the final stage as found in the case of Parribacus japonicus Holthuis, 1960 (Table 2). In addition, Robertson (1969b) compared the dactylus lengths of pereiopods and showed that the dactylus of the Table 2. Comparison of selected larval characteristics in late-stage phyllosomas between arctidine and Parribacus species. Species Stage Outline of cephalic shield TL (mm) CL (mm) CL/CW ratio CW/TW ratio Antenna segment Exopod on P5 Posterior concave margin of thorax References Subfamily Arctidinae Genus Arctides A. guineensis X Ovoid, narrow 29.0 25.2a1.81a0.81a5+Between P5 21 A. regalis VIII Ovoid, narrow 29.5 24.8a1.82a0.79a2+Between P5 22 A. regalis VIII Ovoid, narrow 21.1 17.3 1.73 0.88 2 +Between P5 23 Genus Scyllarides S. herklotsii IXbEllipsoid 19.8 16.0a1.42a1.23a3−Between P4 10 S. squammosus X Ellipsoid 33.4 26.3a1.37a1.18a(5) −Between P4 22 S. squammosus X Ellipsoid 31.0 24.0 1.30 1.07a5−Between P4 18 S. squammosus X Ellipsoid 27.1–31.1 20.2–24.4 1.37–1.41 1.12–1.22 3a−Between P4 20 S. squammosus XI Ellipsoid 36.6–40.0 28.5–30.2 1.32–1.38 1.19–1.23 3a−Between P4 20 S. squammosus XII (final) Ellipsoid 47.0–51.0 43.5a1.31a1.13a5−Between P4 18 S. squammosus XII (final) Ellipsoid 47.0 45.2 1.75 1.24a3 r Between P4 16 S. nodifer XcEllipsoid 26.6 22.9a1.43a1.21a5−Between P4 6 S. sp. VII Ellipsoid 23.6 18.9 1.39 1.18 3 −Between P4 20 S. sp-B X Ellipsoid 27.3 21.8 1.35 1.26 5a−Between P4 20 S. sp-C XII Ellipsoid 48.9 34.0 1.30 1.20 5a−Between P4 20 S. aequinoctialis X Obovoid, wide 28.8 21.2a1.30a1.30a5+Between P5 13 S. astori IXbObovoid, wide 23.4 17.4a1.28a1.28a4a+Between P5 6 S. elisabethae Xb,d Obovoid, wide 22.6a17.5a1.31a1.25a4a+Between P5 8 S. haanii X Ellipsoid 39.5 28.8 1.15 1.40 3 +Between P5 This study Subfamily Ibacinae Genus Parribacus P. antarcticus IX Ellipsoid, wide 33.3–34.0 27.6e1.21e1.24e1+Between P5 24 P. antarcticus IX Ellipsoid, wide 32.4 26.8 1.29 1.22 1 +Between P5 25 P. antarcticus X Ellipsoid, wide 41.8 32.5a1.16a1.27a1+Between P5 22 P. antarcticus X Ellipsoid, wide 50.7 39.8 1.19 1.23 1 +Between P5 25 P. antarcticus XII (final) Obovoid, wide 77.0–80.0 55.8a1.12a1.33a1+Between P5 26 P. japonicus XI Obovoid, wide 70.9 52.8 1.03 1.44 1 +Between P5 25 P. japonicus XI Obovoid, wide 60.8 47.5 1.07 1.43 1 +Between P5 27 P. japonicus (Subfinal) Obovoid, wide 73.0 55.4 1.08 1.45a(2) +Between P5 27 P. japonicus XII (final) Obovoid, wide 80.0 57.0 1.19 1.24a2+Between P5 27 Abbreviations: CL, cephalic shield length; CW, cephalic shield width; P, pereiopod; r, rudiment; TL, total length; TW, thorax width; “+” means present; “−” means absent. a Calculated or judging from the original text figures; b applying Robertson’s (1969a) stage key; c corresponding to Sim’s (1965) stage definition; d as Species Z; e based on the maximum value in their Table 2. References (continued from Table 1): 21 Robertson (1969b); 22 Johnson (1971); 23 Chow et al. (2021); 24 Palero et al. (2014b); 25 Chow et al. (2022b); 26 Prasad et al. (1975); 27 Yoneyama and Takeda (1998). Larval description of Scyllarides haanii 333 pereiopod 4 is larger in Parribacus phyllosoma than in those of the Arctidinae. Phyllosoma specimens from planktonic collections, but not only for the Scyllaridae, are fragile and usually have many missing parts, especially the distal part of the endopod in pereiopods. Thus, all diagnostic characteristics of phyllosomas are completely available in rare cases. It is, therefore, essential to keep this practical matter in mind when selecting morphological characteristics for the classification or identification of phyllosoma larvae. In conclusion, with reference to Sekiguchi’s (1986) key, we tentatively present an updated key to scyllarid phyllosomas from the late to final stages as below. Key to Scyllarid Phyllosmas from the Late to Final Stages 1. Maxilliped 3 with setose exopod Palinuridae (Stridentes)1 — Maxilliped 3 without setose exopod 2 2. Antenna cylindrical, longer than antennule Palinuridae (Silentes)2 — Antenna dorsoventrally compressed, shorter than antennule 3 (Scyllaridae) 3. Cephalic shield bilobed reniform with dorsal carinae and spines Ibacus Leach, 1815 — Cephalic shield rounded ellipsoid, without dorsal ornaments 4 4. Pleon width more than half of thorax 5 — Pleon width less than half of thorax 6 5. Cephalic shield circular or wider than long; telson with posterolateral spines Scyllarinae — Cephalic shield bilobed; telson without posterolateral spines Theninae 6. Pereiopod with subexopodal spine Evibacus Smith, 1869 — Pereiopod without subexopodal spine 7 7. Cephalic shield narrower than thorax Arctides Holthuis, 1960 — Cephalic shield slightly or notably wider than thorax 8 8. Pereiopod 5 without setose exopod Scyllarides Gill, 1898 (in part)3 — Pereiopod 5 with setose exopod 9 9. Antenna unsegmented or two-segmented; dactylus of pereiopod 4 long Parribacus Dana, 1852 — Antenna more than three-segmented; dactylus of pereiopod 4 short Scyllarides (in part)4 1 Panulirus White, 1847, Palinurus Weber, 1795, Justitia Holthuis, 1946, Linuparus White, 1847, etc.; 2 Jasus Parker, 1883, Sagmariasus Holthuis, 1991; 3 S. herklotsii (Herklots, 1851), S. nodifer (Stimpson, 1866), S. squammosus (H. Milne Edwards, 1837); 4 S. aequinoctialis (Lund, 1793), S. astori (Holthuis, 1963), S. elisabethae (Ortmann, 1894), S. haanii (De Haan, 1841). Acknowledgments Our cordial thanks to the captain and crews of R/V Kaiyo-Maru, Fisheries Agency of Japan, for their invaluable support in the research cruise. We are grateful to Dr. M. Sakai for providing us with valuable data on the research cruise. Our thanks are also due to the anonymous reviewers for their valuable suggestions and comments. We also dedicate this paper to the late Dr. Hideo Sekiguchi, Professor Emeritus of Mie University, who passed away in January 2023. Authors Contributions Kooichi Konishi: Conceptualization; Resources; Visualization; Supervision; Writing – original draft. Takashi Yanagimoto: Investigation; Writing – review & editing. Seinen Chow: Funding acquisition; Investigation; Writing – review & editing. Funding This study was partially supported by the project on the Evaluation of the Status of International Fishery Resources by the Fisheries Agency of Japan. Declarations Competing interests. The authors declare no conflicts of interest. References Altshul, S. F., Gish, W., Miller, W., Myers, E. W., and Lipman, D. J. 1990. Basic local alignment search tool. Journal of Molecular Biology 215: 403–410. doi:10.1016/S0022-2836(05)80360-2 Báez, P., Araneda, C., Burns, L., and Navarrete, C. 2022. 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