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Research Article
Verification of two barracudina species Lestrolepis japonica (Tanaka, 1908) and L. philippina (Fowler, 1934) (Aulopiformes, Paralepididae)
expand article infoHsuan-Ching Ho‡§, Toshio Kawai|
‡ Australian Museum, Sydney, Australia
§ National Kaohsiung University of Science and Technology, Kaohsiung, Taiwan
| Hokkaido University, Hakodate, Japan

Corresponding author: Hsuan-Ching Ho ( ogcoho@gmail.com )

Academic editor: Mao-Ying Lee
© 2024 Hsuan-Ching Ho, Toshio Kawai.
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: Ho H-C, Kawai T (2024) Verification of two barracudina species Lestrolepis japonica (Tanaka, 1908) and L. philippina (Fowler, 1934) (Aulopiformes, Paralepididae). In: Ho H-C, Russell B, Hibino Y, Lee M-Y (Eds) Biodiversity and taxonomy of fishes in Taiwan and adjacent waters. ZooKeys 1220: 79-92. https://doi.org/10.3897/zookeys.1220.125860
ZooBank: urn:lsid:zoobank.org:pub:29628E7F-C73B-4ADA-B2C7-2EFD10396909
Open Access

Abstract

Examination of the type series of Lestidium japonica Tanaka reveals that the generally accepted concept of this species does not accord with the type series. A historical review of the literature showed that the characterization of L. japonica changed over time, and what has been recognized as “L. japonica” for more than 70 years actually represents a distinct and different species. Among the junior synonyms of “L. japonica”, Paralepis philippinus Fowler, 1934 is resurrected as a valid species herein in a new combination, Lestrolepis philippina. Lestrolepis nigroventralis Ho, Tsai & Li is recognized as a junior synonym of L. japonica herein. Revised diagnostic characteristics for both L. japonica and L. philippina are provided, along with comments on related names to verify their nomenclatural status.

Key words

Biodiversity, ichthyology, ICZN, nomenclature, taxonomy

Introduction

Naked barracudinas, often classified as the Lestidiini, Lestidiinae, or sometimes Lestidiidae, are a group of small, slender fishes found worldwide from the surface to the deep sea. This group comprises seven genera with approximately 60 species, although some remain problematic (Ho et al. 2019a, 2019b; Ho pers. data). Among the genera, Lestrolepis can be distinguished from all others by having a small light organ (black dot) in front of the orbital margin and a ventral light organ divided into two branches, which are located inside the belly but visible through the translucent muscle. The genus Lestrolepis had few nominal species until recently when Ho and Golani (2019) and Ho et al. (2019a) reviewed and described several new species.

The taxonomic history of Lestrolepis is complex and somewhat confusing, even more so as the author, Robert Rees Harry (Harry 1953a, 1953b), later published under the name Robert R. Rofen (Rofen 1960, 1966). Three species have been commonly recognized in the previous literature, namely Lestrolepis japonica (Tanaka, 1908), Lestrolepis intermedia (Poey, 1868), and Lestrolepis luetkeni (Ege, 1933). However, Ho and Golani (2019) suggested that L. luetkeni belongs to Lestidiops and resurrected Lestrolepis pofi (Harry, 1953a), a name previously considered a junior synonym, as a valid name for the fish formerly referred to as L. luetkeni. Furthermore, Ho et al. (2019a) proposed that L. intermedia is restricted to the Atlantic Ocean and designated the western Pacific population as a new species, Lestrolepis nigroventralis Ho, Tsai & Li, 2019.

Recently, the second author (TK) examined specimens deposited in the fish collection of Hokkaido University and identified inconsistencies between the original description and the specimens currently recognized as L. japonica. Examination of the holotype (Fig. 1) and paratype of L. japonica showed them to be indistinguishable from L. nigroventralis, suggesting that the latter should be regarded as a synonym of L. japonica and that the species previously generally referred to as “L. japonica” requires a new name. The first author (HH) examined the type series of Paralepis philippinus (Fig. 2), previously regarded as a junior synonym of L. japonica, and found them to be identical to what was commonly recognized as L. japonica in the western Pacific Ocean. Additionally, several other names were implicated in the synonymy or misidentifications under these names (Table 1).

Figure 1. 

Holotype of Lestidiun japonicum Tanaka, 1908 A lateral view; arrows point to origins of dorsal fin (DFO), pelvic fin (VFO), and anal fin (AFO) B lateral view of head.

Figure 2. 

Types of Paralepis philippinus (= Lestrolepis philippina) A holotype, USNM 92323, ca 118 mm SL B paratype, USNM 93414, 126 mm SL; arrows point to origins of dorsal fin (DFO), pelvic fin (VFO), and anal fin (AFO).

Table 1.
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Verification of some name records of Lestrolepis and their current status.

Publication Name used Correct names
Poey 1868 Paralepis intermedius Valid as Lestrolepis intermedia
Tanaka 1908 Lestidium japonicum Valid as Lestrolepis japonica
Ege 1933 Paralepis luetkeni Valid as Lestidiops luetkeni
Ege 1933 Paralepis bellottii Synonym of Lestidiops luetkeni
Fowler 1934 Paralepis philippinus Valid as L. philippina
Fowler 1944 Sudis vanderbilti Synonym of L. intermedia
Harry 1953a Lestidium japonicum L. philippina
Ege 1953 Lestidium intermedium (in part) L. japonica
Kamohara 1955 Lestidium japonicum L. japonica
Rofen 1966 Lestrolepis japonica L. philippina
Fujii 1984 Lestrolepis japonica L. philippina
Fujii 1984 Lestrolepis intermedia L. japonica
Nakabo 2000, 2002 Lestrolepis japonica L. philippina
Kim et al. 2007, 2020 Lestrolepis japonica L. philippina
Ho et al. 2019a Lestrolepis nigroventralis Synonym of L. japonica
Ho et al. 2019a Lestrolepis japonica L. philippina

To clarify the identification of these species, we conducted a study on the type series of Lestrolepis japonica and L. philippinna and provide here revised diagnoses or comments for the species related to these names. In the present study we confirm the validity of L. japonica, synonymize L. nigroventralis with L. japonica, resurrect L. philippina, and clarify the taxonomic history of these species.

Materials and methods

Methods for taking morphometrics and meristics followed Ho et al. (2019a). Most morphometric and meristic data are adopted from Ho et al. (2019a), with those of additional specimens examined.

Abbreviations

SL, standard length; HL, head length; TL, total length; VFO, DFO, AFL, the origins of pelvic fin, dorsal fin and anal fin, respectively; V–D, space between VFO and DFO; V–A, space between VFO and AFO. Other abbreviations followed Ho and Lin (2023).

Results

Genus Lestrolepis Harry, 1953

On the taxonomic issues

Lestidium japonicum was described based on two types collected from Sagami Bay (ZMUT 2013 and 2014). In the synonymy of this species, Harry (1953a) included Paralepis bellottii Ege, 1933 as a queried synonym and Paralepis philippinus Fowler, 1934 as a junior synonym of L. japonicum. Subsequently, Harry (1953b) re-assigned japonicum to Lestrolepis, and this placement in Lestrolepis has been widely accepted by subsequent authors.

Post (1972) suggested that L. japonica “possibly is a synonym of Lestrolepis intermedia Poey, 1868". Examination of the holotype of L. japonica revealed that it is indeed similar to L. intermedia but that it differs in several characters. Consequently, specimens recognized as “Lestrolepis japonica” in much of the literature belong to another species that requires a name, and the following two names listed as synonyms of “japonicum” by Harry (1953a) should be considered as candidates for this species.

The original description of Paralepis bellottii was based on a single specimen (60 mm SL, 64 mm TL). This species was synonymized with Lestidium nudum Gilbert, 1905 and Paraelpis luetkeni Ege, 1933 by Ege (1953:52). Rofen (1960: 206) synonymized the name with “Lestidium leutkeni” and Rofen (1966) later placed it in the genus Lestrolepis. The first author examined the holotype of bellottii and found it to be very similar to the holotype of luetkeni. Type specimens of both are also different from Lestidium nudum, rejecting the synonymy proposed by Ege (1953). In fact, luetkeni is placed in the genus Lestidiops by Ho and Golani (2019), and the status of bellottii will be discussed in another work prepared by HH.

The original description of Paralepis philippinus was based on eight type specimens collected from the Philippines. Examination of the type series revealed that it is different from that of L. japonica and is what has been commonly recognized as “L. japonica” in the western Pacific Ocean.

Consequently, neither philippinus nor leutkeni or bellottii, can be regarded as synonyms of japonica, but can be considered here to represent what has been called “L. japonica” in the western Pacific (Table 1). The oldest name, philippina is herein resurrected as valid in a new combination Lestrolepis philippina (Fowler, 1934) for this species.

Ho et al. (2019a) described Lestrolepis nigroventralis to accommodate the population of what has been called “Lestrolepis intermedia” in the western Pacific Ocean, i.e. Japan, Korea, and Taiwan, etc. As the type series is the same as that of L. japonica (Table 1), this name is now recognized as a junior synonym of L. japonica. In addition, the specimens recognized as L. japonica by Ho et al. (2019a) are now re-identified as L. philippina.

Records of Lestrolepis japonica and L. intermedia in the western Pacific

As mentioned above, Harry (1953a) mistakenly considered Lestrolepis philippina as a junior synonym of L. japonica without examining the vertebral counts of the type series of L. japonica. This oversight led to a change in the definition of L. japonica based on the type series of L. philippina, resulting in subsequent effects. For instance, Rofen (1966: 381) differentiated L. intermedia from L. japonica based on different vertebral numbers (91–98, vs 84–89), indicating that his L. japonica was actually L. philippina. Subsequent literature records of L. japonica following Harry’s definition recognized those with fewer vertebrae as L. japonica (see synonymy below).

The earliest records of Lestrolepis intermedia in the Western Pacific Ocean are found in Ege (1953), who documented juveniles of L. intermedia collected from various locations worldwide, including Japan, Taiwan, and the Philippines (also see Rofen 1966). However, subsequent publications in the western Pacific Ocean (e.g., Matsubara 1955) did not evidently document this name.

Fujii in Masuda et al. (1984:77) provided a brief description of Lestrolepis intermedia and mentioned that “L. intermedia and L. japonica are often collected by shrimp-trawl net in Suruga Bay”. Fujii also provided total vertebrae counts of 95–97 and total lateral-line scales of 74–81 for his L. intermedia, and 84–87 and 62–69, respectively, for his L. japonica. These definitions for both species evidently were followed by all subsequent publications in Japan and nearby areas (e.g., Nakabo 1993, 2000, 2002; Shen et al. 1993, Kim et al. 2007), while Ho et al. (2019a) went further and mistakenly recognized the western Pacific population of L. intermedia as a new species, L. nigroventralis.

Lestrolepis japonica (Tanaka, 1908)

Figs 1, 3, Table 2

Lestidium japonicum Tanaka, 1908:27 (type locality: Sagami Sea, Japan); Jordan et al. 1913: 50 (Sagami Sea, Japan; list); Okada and Matsubara 1938: 61 (Sagami Sea, Japan; key); Matsubara 1941: 8 (in part: Japan); Matsubara 1955: 262 (in part: Sagami Sea, Wakayama and Mie, Suruga Bay, Japan; key); Matsubara 1963: 262 (in part: Sagami Sea, Wakayama and Mie, Suruga Bay, Japan; key).

Lestrolepis intermedia (non Poey, 1868): Fujii in Masuda et al. 1984:77 (Japan; short description); Chen and Yu 1986:324 (Taiwan; synopsis); Nakabo 1993:319 (Japan, picture key); Shinohara et al. 1996:164 (Honshu; list); Nakabo 2000:371 (Japan, picture key); Nakabo 2002:371 (Japan, picture key); Shinohara et al. 2005:409 (Ryukus Is.; list); Kim et al. 2007:63 (Korea; new record); Ikeda and Nakabo 2015: 312 (Japan; short description).

Lestrolepis nigroventralis Ho, Tsai & Li, 2019:123 (type locality: off Dong-gang, Pingtung, southwestern Taiwan).

Status of the holotype

Tanaka (1908) indicated that the holotype of his Lestidium japonicum was registered as 2013 (now ZMUT 2013) with a length of 174 mm SL, and the paratype (in the Table, without a catalog number) was 190 mm SL. Regardless, Harry (1953a) gave the holotype as ZMUT 2014, and the paratype as ZMUT 2013. Notwithstanding, Post (1972) documented the types and recognized ZMUT 2013 as the holotype and ZMUT 2014 as the paratype. We examined both types and re-measured the holotype (ZMUT 2013) as 159.5 mm SL and the paratype (ZMUT 2014) as approximately 180 mm SL, the latter being in poor condition.

Figure 3. 

Fresh condition of Lestrolepis japonica (Tanaka, 1908), NMMB-P027930, 244 mm SL A lateral view B lateral view of head C lateral view of body showing origin of dorsal fin (DFO), pelvic fin (VFO), and anal fin (AFO) D lateral view of caudal region.

Synonym name

Lestrolepis nigroventralis was described to accommodate the Pacific population of L. intermedia. However, because of its nearly identical morphology (Table 2), it is now recognized here as a junior synonym of L. japonica. The following data are mainly derived from types and non-types of L. nigroventralis, combined with the type series examined by us.

Table 2.
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Selected morphological and meristic data of L. japonica and L. philippina. * Data of “L. nigroventralis” and ** of “L. japonica” taken from Ho et al. (2019a). HT = holotype; PT = paratype(s).

L. japonica L. philippina
Types Types of L. nigroventralis Types Non-types**
HT PT HT Types* HT PT
SL (mm) 159.5 ca 180 230 180–249 (n = 11) 118.2 109.6–128.0 (n = 4) 88.0–222 (n = 29)
Proportion (%) Mean (Range) Mean (Range)
HL/SL 20.9 21.1 20.0 19.6 (18.9–20.7) 20.8 21.3–22.0 20.5 (18.0–21.8)
BD/SL 5.7 5.2 5.3 5.2 (4.9–5.6) 8.5 7.1–7.9 6.9 (5.9–8.6)
PreD/SL 63.7 – 62.8 62.7 (61.7–63.3) 58.8 59.4–60.9 61.8 (60.2–63.5)
PreV/SL 52.7 – 52.1 51.6 (50.6–52.6) 51.2 51.6–53.4 53.0 (52.3–55.2)
PreA/SL 73.3 – 73.5 73.3 (71.9–74.8) 72.8 73.0–75.2 74.9 (75.8–76.6)
ED/HL 20.1 17.9 15.5 16.1 (14.8–17.3) 19.1 18.1–19.1 18.4 (15.9–22.6)
SN/HL 54.7 53.8 52.4 54.4 (52.4–57.0) 52.8 49.8–52.8 50.8 (47.7–53.4)
HD/HL 27.1 24.8 26.3 26.4 (25.4–27.4) 30.9 27.8–30.8 28.5 (28.6–28.5)
UJ/HL 51.4 49.6 49.1 49.0 (47.2–50.4) 47.6 47.7–49.5 46.8 (44.0–50.0)
V–D/V–A 53.1 51.0 50.0 51.0 (47.6–55.0) 35.3 34.0–36.1 39.1 (32.5–43.1)
Meristics
Anal-fin rays 42 ca 40 42 41–43 ca 38 37–38 36–40
Vertebrae
PHV 34 33 33 32–35 29 30–31 29–32
PVV 33 34 34 33–35 30 30–32 30–33
PDV 43 44 44 43–46 38 38–39 36–40
PAV 53 55 55 53–57 48 49–51 48–51
CV 60 61 62 60–66 55 56–59 52–58
TV 94 95 95 94–98 85 86–89 84–88
V–D 9 10 10 9–11 8 7–8 5–9
Lateral-line scales
PVLL 34 – 34 33–36 30 30–32 31–32
PDLL 44 – 44 43–46 38 39–40 38–40
PALL 54 – 54 53–57 49 49–51 49–50
TLL ca 81 – 78 75–81 71 68–70 65–72

Distinguishing features

A species of Lestrolepis with DFO situated at about midline of V–A, V–D 46.8–55.0% of V–A; anal-fin rays 41–43 (mainly 41–42); lateral-line scales: PVLL 33–36 (34–35), PDLL 43–46 (44–46), PALL 53–57 (55–56), TLL 75–81 (77–80); vertebral counts: PHV 32–35 (mainly 32–34), PVV 33–35 (34–35), PDV 43–46 (44–45), PAV 53–57 (54–55), CV 60–66 (61–64), TV 94–98 (94–97); vertebrae between DFO and VFO 9–11. Body slender, body depth at pectoral-fin base 15–19 times in SL; band of melanophores along abdominal margin. Attains 249 mm SL.

Distribution

Known from the northwestern Pacific Ocean off Japan, Taiwan, and Korea. Records from other localities require verification.

Remarks

The taxonomic concept of what has long been called Lestrolepis japonica is now revised based on the type series and additional specimens. A review of the literature with documentation of L. japonica is listed in the synonymy above, although there might be more records that are not included therein. Detailed comparisons with congeners are provided in Ho et al. (2019a, as L. nigroventralis).

There are some minor differences found in the types compared to the non-types (Table 2). The head length is slightly larger (20.9–21.1%, vs 18.9–20.7% SL); the predorsal length is slightly larger in the holotype (63.7%, vs 61.7–63.3% SL; not available for the paratype); the eye diameter is slightly larger (17.9–20.1%, vs 14.8–17.3% HL); the upper jaw of the holotype is slightly longer (51.4%, vs 47.2–50.4% HL). These proportional measurements may be attributed either to population variation or due to long-term preservation of the types. The meristic values of types fall entirely within the range of non-types (Table 2).

Lestrolepis philippina Fowler, 1934

Figs 2, 4, Table 2

Paralepis philippinus Fowler, 1934: 281, fig. 42 (type locality: Varadero Harbor, Philippines).

Lestidium japonicum (not of Tanaka): Matsubara 1941: 8 (in part: Japan); Matsubara 1955: 262 (in part: Sagami Sea, Wakayama and Mie, Suruga Bay, Japan; key); Matsubara 1963: 262 (in part: Sagami Sea, Wakayama and Mie, Suruga Bay, Japan; key).

Lestrolepis japonica (not of Tanaka): Fujii in Masuda et al. 1984:77 (Japan; short description); Nakabo 2000:371 (Japan; picture key); Paxton in Randall and Lim 2000:592; Nakabo 2002:371 (Japan; key); Shinohara et al. 2005:409 (Honshu; list); Kim et al. 2007:64 (Korea; new record); Ikeda and Nakabo 2015: 312 (Japan; short description); Motomura et al. 2017:51; Ho and Golani 2019:578 (mentioned); Ho et al. 2019a:127 (Taiwan and Japan; description); Kim et al. 2020:67; Misawa et al. 2020:273.

Taxonomy

Fowler (1934) originally described Lestrolepis philippina from the Philippines. Subsequently, Harry (1953a) synonymized L. philippina with L. japonica, providing a description based on the type series of L. japonica, L. philippina, and additional specimens. Although the name L. philippina was not mentioned in subsequent literature, it is likely that many records recognized as L. japonica are, in fact, misidentifications of L. philippina. In a more recent study, Ho et al. (2019a) presented a detailed description of what was previously identified as L. japonica, now re-identified as L. philippina.

Figure 4. 

Fresh condition of Lestrolepis philippina (Fowler, 1934), NMMB-P027934 (1 of 17), ca 175 mm SL A lateral view B lateral view of head C lateral view of body showing origin of dorsal fin (DFO), pelvic fin (VFO), and anal fin (AFO) D lateral view of caudal region.

Distinguished features

A species of Lestrolepis with DFO situated well before midline of V–A, V–D 32.5–43.1% of V–A; anal-fin rays 36–40 (mainly 37–39); lateral-line scales: PVLL 30–32 (31), PDLL 38–40 (38–39), PALL 49–51, TLL 65–72 (65–68); vertebral counts: PHV 29–32, PVV 30–33 (30–32), PDV 36–40 (37–40), PAV 48–51, CV 52–58 (54–57), TV 82–88 (84–88); vertebrae between DFO and VFO 5–9 (7–8). Body moderately slender, body depth at pectoral-fin base 13–16 times in SL; narrow band of melanophores along abdominal margin. Attains 222 mm SL.

Distribution

Widespread in the western Pacific Ocean, with confirmed records from Japan, Taiwan, the Philippines, northwestern Australia, and the South China Sea.

Remarks

Some minor differences are observed between the holotype and non-types (Table 2). The predorsal length (58.8%, vs 60.2–63.5% SL), preventral length (51.2%, vs 52.3–55.2% SL), and preanal length (72.8%, vs 75.8–76.6% SL) are slightly smaller in the holotype compared to paratypes and non-types. The head depth is slightly larger (30.9% SL) compared to that of paratypes and non-types (27.8–30.8% SL). These differences may be attributed to individual variation or long-term preservation effects. The meristic values of the type series fall well within those of the non-types (Table 2).

Discussion

Several ambiguities in earlier studies have led to subsequent problems, including the inadvertent naming of a new species. Tanaka (1908) described Lestidium japonicum with 42 anal-fin rays in the holotype and 49 in the paratype. Harry (1953a) counted 42 and 41, respectively, for the same individual (note that Harry switched the holotype and paratype). Jordan et al. (1913) and Okada and Matsubara (1938) included Lestidium japonicum in their species list as known from Sagami Sea. On the other hand, Hubbs (1916) cited Tanaka (1908) and gave 42–49 anal rays. Parr (1928) noted that Lestidium japponicum [sic] is only known from the coast of Japan.

Matsubara (1941) reported two specimens (178.8 mm SL and 102.2 mm SL) of this species collected from Suruga Bay, Japan, the former specimen having 43 anal-fin rays, and the latter 35. Harry (1953a: 187) suggested the first one is Lestidium japonicum, whereas the latter was unknown. We concur that the former is Lestrolepis japonica, and the latter is L. philippina based on their counts of anal-fin rays (cf. 40–43 in L. japonicus and 36–40 in L. philippina; Table 2). Matsubara (1941) also mentioned that japonicum closely resembles L. philippinus. This could have been the beginning of confusion of the species (see also Matsubara 1955, 1963).

Harry (1953a) reported examining the types and additional specimens of L. japonica and identified L. philippina as a junior synonym. However, given clear differences, such as the position of DFO, the relatively slender body, snout, and jaws, as well as different vertebral and lateral-line counts, it remains uncertain why Harry considered them senior synonyms of L. philippina, perhaps being influenced by Matsubara (1941). It seems likely that Harry never examined radiographs of the types of L. japonica; otherwise, he might have noticed the distinctly high vertebral counts in these two specimens.

Ege (1953) documented juveniles of Lestrolepis intermedia (as Lestidium intermedium) from the western Pacific, i.e. Japan, Taiwan, and the Philippines, etc. Assuming he identified these specimens correctly, for example, regarding the position of DFO, his specimens should be L. japonica in this work. However, he never considered the names “japonicum” or “philippinus” in his publication. Based on the museum collection (Ho pers. obs.), L. philippina is far more abundant compared to L. japonica, and it remains unknown why Ege did not recognize such a distinct form. It is notable that Ege (1953) recognized from this region Lestidium nudum, which has similar fin position. It is likely Ege misidentified L. philippina as Lestidium nudum, because the latter is more or less restricted to the Hawaiian Islands and the central Pacific (Ho pers. data).

Fujii in Masuda et al. (1984) appears to have followed Harry’s (1953b) definition and recognized the population with few vertebrae as L. japonica, which is now confirmed to be a misidentification of L. philippina. He also identified L. intermedia from Japan and provided a short description. His provided counts of 95–97 total vertebrae and 74–81 lateral-line scales indicate that his description was, in fact, that of L. japonica.

Regarding other paralepidid species, some taxonomic problems have been reviewed, such as the resurrection of Lestrolepis pofi and the placement of L. leutkeni in Lestidiops (Ho and Golani 2019). Matching adults to these species with only juveniles known, or redescriptions of species have been undertaken (Ho and Huang 2022a, 2022b; Ho and Lin 2023; Ho and Tsai 2023). However, there are several species with unknown status that still require further investigation (Ho pers. data).

This work provides an opportunity to underscore the importance of not relying solely on previously published works but also examining type material in museum collections, even for well-known species.

Material examined

Lestrolepis japonica : Holotype. ZMUT 2013 (159.5), Sagami Bay, Japan. Paratype. ZMUT 2014 (ca. 180), same as holotype. Non-types. HUMZ 71212 (1, 183), Fukushima, Japan, trawl net, 200–300 m, 8 Nov. 1977. HUMZ 71248 (1, 243), Hayakawa Port, Odawara, Kanagawa, Japan, 7 Nov. 1977. HUMZ 71844 (1, 193), HUMZ 71845 (1, 296), HUMZ 71846 (1, 184), HUMZ 71847 (1, 213), HUMZ 71848 (1, 186), Hayakawa Port, Odawara, Kanagawa, Japan, 5 Nov. 1977. HUMZ 134997 (1), 38°35'30.0"N 141°59'48.0"E, off Minamisanriku, Miyagi, Japan, bottom trawl, 331 m, 17 Oct. 1994. HUMZ 163755 (1, 120), 38°24'32.4"N, 142°04'13.2"E Miyagi, 461 m, 12 Oct. 1999. HUMZ 163832 (1, 123), HUMZ 163833 (1, 125), Miyagi, 38°23'09.6"N, 142°07'27.0"E, 552 m, 26 Oct. 1999. HUMZ 206777 (1, 162), 38°24'57"N, 142°02'50.4"E, Miyagi, Japan, 425–430 m, 30 Oct. 2009. HUMZ 206833 (1, 140), 38°22'13.2"N, 142°03'13.8"E, Miyagi, bottom trawl, 412 m, 9 Oct. 2009. HUMZ 209243 (1, 92), 39°35'24.6"N, 142°31'E, 890-897 m, 7 Oct. 2010. HUMZ 214579 (1, 134), 39°05'15.6"N, 142°09'34.8"E, off Tohoku, 398–412 m, 19 Oct. 2011. HUMZ 222513 (1, 183), Miyagi, 38°52'01.2"N, 142°04'24.6"E, 357 m, bottom trawl, 23 Oct. 2013. HUMZ 226859 (1, 155), 39°01'59.4"N, 142°12'55.8"E, off Kesennuma, Miyagi, Japan, bottom trawl, 575 m, 23 Oct. 2015. HUMZ 226876 (1, 146), HUMZ 226877 (3), 37°36'56.4"N, 141°50'31.8"E, off Souma, Fukushima, Japan, 380 m, 5 Nov. 2015. HUMZ 226942 (2), 37°43'N, 141°53'51"E, off Souma, Fukushima, Japan, bottom trawl, 411 m, 29 Oct. 2015. NSMT-P13816 (2, 147–154), Suruga Bay, Shizuoka, Japan, 17 Nov. 1968. NSMT-P48916 (1, 168), south of Sagami Bay, Japan, 4 Nov. 1995. NSMT-P48931 (1, 154), off Tohoku, northern Japan, trawl, 5 Nov. 1995. NSMT-P58786 (3, 75–130), 39°00'36"N, 143°32'06"E, off Pacific coast between Miyagi Pref. and Iwate, 550–578 m, Japan, mesopelagic trawl, 29 Jul. 1996. NSMT-P 58787 (1, 100), 39°00'36"N, 143°32'06"E, between Miyagi Pref. and Iwate, 650–677 m, mesopelagic trawl, 29 Nov. 1996. NSMT-P58788 (1, 115), NSMT-P58789 (1, 93), 39°02'24"N, 143°30'07.2"E, between Miyagi Pref. and Iwate, northern Japan, 650–677 m, mesopelagic trawl, 30 Jul. 1996. NSMT-P65464 (1, 176), 37°45'54"N, 142°09'32"E, off Fukushima, northern Japan, 647–676 m, otter trawl, 19 Oct. 2002. NSMT-P65466 (1, 136), 36°53'45.6"N, 141°33'43.2"E, off Ibaraki, Japan, 495–530 m, otter trawl, 20 Oct. 2002. NSMT-P67563 (1, 158), 31°20'39.1"N, 128°10'53"E, southern Japan, 392 m, otter Trawl, 8 Nov. 2003. NSMT-P67589 (1, 237), 28°59'47"N, 127°09'21.2"E, Ryukyus, 350 m, otter Trawl, 5 Nov. 2003. NSMT-P91547 (1, 170), 38°21'32.4"N, 141°56'24.0"E, off Miyagi, Japan, 280 m, trawl, 3 Oct. 2007. NSMT-P102802 (1, 169), 36°58'26.4"N, 141°25'43.3"E, off Fukushima, Japan, 251-252 m, otter trawl, 26 Oct. 2006. Also listed as Lestrolepis nigroventralis in Ho et al. (2019a), including the type series.

Lestrolepis philippina : Holotype. USNM 92323 (118.2), Varadero Bay, southern Luzon, Mindoro, Philippine, 22 Jul. 1908. Paratypes. USNM 93414 (2, 109.6–125), Noble Point, Tulayan Island, Sulu, Philippine, 283 m, 15 Sep. 1909. CAS-SU 14970 (1), USNM 135257 (2, 121–128), Anchorage, Dupon Bay, Leyte, Philippine, 17 Mar. 1909. Non-types. HUMZ 146645 (1, 228), 25°37'16.8"N, 126°05'21"E, Miyoko, Okinawa, 388–394 m, 2 Aug. 1994. HUMZ 222365 (1, 153), HUMZ 222366 (1, 165), HUMZ 222367 (1, 180), HUMZ 222368 (1, 185), HUMZ 222462 (1, 144), Dong-gang, Taiwan, 7 Nov. 2013. NSMT-P59709 (2, 87–96), 35°50'38.4"N, 152°59'49.2"E, 0–20 m, trawl, 24 May 1995. NSMT-P67590 (2, 185-232), 28°59'47"N, 127°09'21.2"E, East China Sea, 350 m, otter Trawl, 5 Nov. 2003. NSMT-P94628 (8, 123–160), Sagami Bay, southern Japan, set net, Tokyo Sea Life Park, 15 Nov. 1988. NSMT-P115090 (1, 138.2), NSMT-P115091 (1, 154.5), Dong-gang fishing port, Taiwan, 17 Oct. 2013, G. Shinohara et al. NSMT-P115637 (1, 142), Dong-gang fishing port, Taiwan, 19 Oct. 2013. NSMT-P128174 (1, 268.8), off Amami-Oshima Is., Ryukyus, Japan, Yamakawa, 26 Jun. 1972. NSMT-P128175 (1, 226.1), off Amami-Oshima Is., Ryukyus, Japan, 26 Jun. 1972. Also listed as L. japonica in Ho et al. (2019a)

Acknowledgements

We thank H. Wada (The University Museum, The University of Tokyo), S.L. Ng (Department of Environmental Biology and Fisheries Science, National Taiwan Ocean University), K. Bemis, A. Reft (NOAA National Systematics Lab), and D. Pitassy (Smithsonian National Museum of Natural History) for providing various assistance, S. Nihei for taking photos of ZUMT specimens, and B. Russell for giving the critical review. This study is supported by the National Kaohsiung University of Science and Technology, Kaohsiung, Taiwan, and JSPS KAKENHI grant number JP23K05364 to TK.

Additional information

Conflict of interest

The authors have declared that no competing interests exist.

Ethical statement

Only museum collection specimens were used in present study.

Funding

This study was supported by the National Science and Technology Council, Taiwan, to HCH, and JSPS KAKENHI Grant (no. JP23K05364) to TK.

Author contributions

H.-C. Ho and T. Kawai took data, photos, and composed and reviewed the manuscript. All authors approved the manuscript.

Author ORCIDs

Hsuan-Ching Ho https://orcid.org/0000-0003-1154-601X

Data availability

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

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