Research Article |
Corresponding author: Keita Koeda ( koeda@sci.u-ryukyu.ac.jp ) Academic editor: Hsuan-Ching HO
© 2024 Keita Koeda, Manabu Bessho-Uehara.
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:
Koeda K, Bessho-Uehara M (2024) A review of the genus Pempheris (Teleostei, Pempheridae) found in Japan and Taiwan. In: Ho H-C, Russell B, Hibino Y, Lee M-Y (Eds) Biodiversity and taxonomy of fishes in Taiwan and adjacent waters. ZooKeys 1220: 123-163. https://doi.org/10.3897/zookeys.1220.126762
|
Species of the genus Pempheris Cuvier, 1829 (Pempheridae) from Japan and Taiwan are taxonomically reviewed based on morphology supported by molecular phylogenetic analysis. Ten species are recognized from these countries: Pempheris adusta Bleeker, 1855, Pempheris familia Koeda & Motomura, 2017, Pempheris japonica Döderlein, 1883, Pempheris nyctereutes Jordan & Evermann, 1902, Pempheris oualensis Cuvier, 1831, Pempheris sasakii Jordan & Hubbs, 1925, Pempheris schwenkii Bleeker, 1877, Pempheris ufuagari Koeda, Yoshino & Tachihara, 2013, Pempheris vanicolensis Cuvier, 1831, Pempheris xanthoptera Tominaga, 1963. Nine of them are distributed in Japan, and five of them in Taiwan. Pempheris sasakii and P. xanthoptera, nominal species that have been regarded as invalid are revalidated, redescribed with diagnoses based on examinations of the holotypes and the specimens collected from Japan. Pempheris sasakii is morphologically similar to P. nyctereutes and has been thought to be a senior synonym of the latter, but the comparison of the holotypes and non-types of both species revealed that the former species is distinguishable from the latter species in having fewer counts of body scales, also genetically supported with a 3.1% mitochondrial DNA sequence divergence. Pempheris xanthoptera is similar to P. schwenkii, but the coloration of their caudal fins is different, and the genetic analysis supported the difference. The distributions of all species of the genus Pempheris in Japanese waters are also described, based on the specimen localities from literature and new material herein.
Distribution, morphology, Pempheris sasakii, Pempheris xanthoptera, sweepers, taxonomy
The family Pempheridae, also known as sweepers, is a group of nocturnal fish widely distributed in the Indo-Pacific and western Atlantic Ocean. This family is currently divided into two genera: Parapriacanthus Steindachner, 1870 and Pempheris Cuvier, 1829. The latter is characterized by having anal-fin soft rays numbering 30–45, the anal-fin base covered with scales and longer than 40% of the standard length, the lateral line extending onto the posterior margin of the caudal fin, and the first interhaemal angled toward the posterior end of the dorsal-fin base (
The taxonomy of the family Pempheridae in Japanese waters was reviewed by
Preserved materials examined in the present study including large numbers of specimens collected by KK are listed in Suppl. material
The nucleotide sequences of mitochondrial 16S ribosomal RNA (16S) and cytochrome oxidase I (COI) were analyzed to infer phylogenetic relationships. DNA was extracted from ethanol-fixed specimens and the sequences of 16S and COI were obtained as described by
Pempheris Cuvier, 1829: 195 [type species: Pempheris touea = Kurtus argenteus Bloch & Schneider, 1801 = Sparus ? compressus (Shaw, 1790): junior synonym of Pempheris compressa (Shaw, 1790)].
Priacanthopsis Fowler, 1906: 122 (type species: Pempheris mulleri: junior synonym of Pempheris schomburgki Müller & Troschel, 1848)].
Catalufa Snyder, 1911: 528 (type species: Catalufa umbra: junior synonym of Pempheris japonica Döderlein, 1883).
Liopempheris Ogilby, 1913: 61 (type species: Pempheris multiradiatus Klunzinger, 1879).
Body shape oval, strongly compressed laterally; body deep, deepest at or near origin of dorsal fin; dorsal outline of head nearly straight or generally curved from snout to origin of dorsal fin; ventral outline of body generally curved to origin of pelvic fin; body depth rapidly decreases at posterior half of body; depth of caudal peduncle < 1/4 of maximum body depth.
Eye large; snout very short; interorbital space slightly convex or flat; two nostrils located just anterior to anterior margin of eye. Mouth large, strongly oblique; lower jaw slightly project beyond upper jaw; villiform teeth on jaws; tip of tongue free from floor of mouth. Lips thin. Gill opening large; outer margin of opercle and preopercle smooth. Gill membranes on left and right sides separate, free from isthmus. Gill rakers long, 6–13 (upper)+17–28 (lower) on first gill arch.
Body and head almost fully covered by strongly or weakly ctenoid scales except for lips and anterior to eye; ~ 1/3 of basal part of anal fin covered with small scales. Lateral line starts from uppermost position of opercle, generally follows dorsal outer margin, through middle of caudal peduncle, and extending to middle of posterior end of caudal fin. Anus slit-like, located just anterior to anal fin. Light organ present in some species (absent in species distributed in the Northern Hemisphere).
Vertebral counts 10+15 (abdominal + caudal), very rarely 10+16; predorsal interneurals 3; 4th interneural supporting 1st dorsal-fin spine, inserted between 2nd and 3rd vertebrae; last interneural with last dorsal ray inserted between neural spines of 11th and 12th vertebrae, or 12th and 13th. First interhaemal supporting 1st and 2nd anal-fin spine, inserted in front of haemal spine of 11th vertebra, and pointing to posterior end of dorsal fin.
Dorsal fin single, triangular; its base short, shorter than longest ray; 5–7 spines, last longest; 8–13 soft rays, 1st or 2nd longest, rapidly shorter posteriorly. Anal fin low, its base very long, length longer than body depth; three spines, last longest; 24–49 soft rays, 1st longest, gradually shorter posteriorly. Pectoral fin pointed posterodorsally; 15–20 rays, uppermost two rays unbranched, 3rd or 4th ray longest, shorter in lower; pectoral-fin length longer than length of longest dorsal-fin ray. Pelvic fin small, with one spine and five soft rays, 1st longest; last ray not connected to body with membrane. Caudal fin triangular, weakly forked.
Body color uniformly silver, copper, or golden without distinct patterns except for Pempheris ornata Mooi & Jubb, 1996 (not in the Northern Hemisphere) which has longitudinal golden stripes on body laterally.
Indo-Pacific Ocean: north to southern Japan, east to Easter Island (not including Hawaii Islands), south to Tasmania, west to South Africa, and the Red Sea (some species migrated from the Red Sea to the eastern part of Mediterranean); western Atlantic Ocean: north to Florida; south to Brazil (
This genus includes a large number of species, and the counts, measurements, and colorations are not very informative in distinguishing them from each other because of the interspecific uniformity and the intraspecific diversity of the results. This had led to significant taxonomic confusions, and the recent jumbled descriptions of abnormally high numbers of new species reported from the Indian Ocean have caused further misunderstandings in the taxonomy of the genus.
Molecular phylogenetic analysis using three methods (ML, NJ, and BI) showed consistent topology, except for the placement of P. ufuagari. In the ML and BI trees, P. ufuagari is positioned as a sister to a clade composed of P. vanicolensis and P. oualensis with low node supporting values. In contrast, the NJ tree places P. ufuagari as a sister to a clade composed of P. vanicolensis and P. adusta. All species analyzed in this study displayed monophyly with high supporting values on their respective nodes.
1 | 12–15 scale rows above lateral line; scales on lateral body strongly ctenoid and adherent, with distinct basal and distal portions ( |
2 |
– | 3½–10½ scale rows above lateral line; scales on lateral body weakly ctenoid and deciduous ( |
3 |
2 | 69–82 pored lateral-line scales; 12 or 13 scale rows above lateral line; 40–44 predorsal scales; 22–24 circumpeduncular scales; blackish blotch on the pectora-fin base absent or faint | P. japonica (southern Japan, Izu Islands, western Japan Sea, rarely in Ryukyu Archipelago; southern Korea) |
– | 84–88 pored lateral-line scales; 14 or 15 scale rows above lateral line; 50–55 predorsal scales; 26 circumpeduncular scales; distinct pupil-sized blackish blotch on the pectoral-fin base present | P. familia (Ogasawara Islands) |
3 | 67–81 pored lateral-line scales; 8½–9½ scale rows above lateral line; 19–27 scale rows below lateral line; snout pointed; scales on ventral and pored lateral-line scales strongly ctenoid; body brownish with golden reflection; lateral line distinctly whitish | 4 |
– | 44–71 pored lateral-line scales; 3½–7½ scale rows above lateral line; 10–18 scale rows below lateral line; snout weakly pointed or rounded; scales on ventral and pored lateral-line scales weakly ctenoid; body pale brown to grey with golden, silver, or copper reflections; lateral line same as uniform color of body | 5 |
4 | 67–77 pored lateral lateral-line scales, usually fewer than 73; 19–22 scale rows below lateral line; body brown, with golden reflection in fresh specimen | P. sasakii (southern Japan, northern Ryukyu Archipelago) |
– | 72–81 pored lateral lateral-line scales, usually > 79; 23–28 scale rows below lateral line; body silver to dark brown in fresh specimen | P. nyctereutes (Taiwan; Vietnam) |
5 | 44–65 pored lateral lateral-line scales; 3½–6½ scale rows above lateral line; black blotch on pectoral-fin base absent; pectoral fin uniformly pink or bright yellow | 6 |
– | 51–71 pored lateral lateral-line scales; 4½–7½ scale rows above lateral line; black blotch on pectoral fin-base present; pectoral fin uniformly pink or upper half dusky | 8 |
6 | 57–65 pored lateral lateral-line scales; 5½–6½ scale rows above lateral line; 12–15 scale rows below lateral line; body with silver reflection; pectoral fin bright yellow; outer margin of anal fin distinctly blackish | P. vanicolensis (Ryukyu Archipelago, Taiwan; western Pacific) |
– | 44–54 pored lateral lateral-line scales; 3½–4½ (usually 3½) scale rows above lateral line; 10–12 scale rows below lateral line; body with golden or silver reflection; pectoral fin pink; outer margin of anal fin faint blackish or translucent | 7 |
7 | Posterior nostril usually oval, rounded; caudal fin bright yellow | P. xanthoptera (southern Japan, Izu Islands, western Japan Sea, northern Ryukyu Archipelago, Ogasawara Islands; southern Korea) |
– | Posterior nostril usually slit-like; caudal fin pink to brown | P. schwenkii (southern Kyusyu, Ryukyu Archipelago; Taiwan; western Pacific) |
8 | 51–62 pored lateral lateral-line scales; 4½–5½ scale rows above lateral line; black blotch on pectoral-fin base faint, usually on posterior 2/3 of its base | P. adusta (southern Japan, Izu Islands, Ryukyu Archipelago, Daito Islands, Ogasawara Islands; Taiwan; western Pacific) |
– | 60–71 pored lateral lateral-line scales; 6½–7½ scale rows above lateral line; distinct black blotch covering entire pectoral-fin base present | 9 |
9 | Usually 7½ scale rows above lateral line; tooth band absent at outside of lips; dorsal and caudal fins bright yellow; pectoral fin uniformly pink; tip of dorsal fin black, but anterior margin not black; outer margin of anal fin distinctly blackish | P. ufuagari (Daito Islands, Ogasawara Islands) |
– | Usually 6½ scale rows above lateral line; tooth band present at outside of lips (in large individuals); dorsal and caudal fins brown; upper half of pectoral fin dusky; anterior margin to tip of dorsal fin black; outer margin of anal fin without black coloration | P. oualensis (Ryukyu Archipelago, Daito Islands, Ogasawara Islands; Taiwan; Western Pacific) |
Pempheris adusta
Bleeker, 1877: 50, pl. 383, fig. 1 (type locality: Ambon Island, Molucca Islands, Indonesia);
Pempheris mangula
(not Cuvier, 1829):
Pempheris oualensis
(not Cuvier, 1831):
Pempheris
sp.:
Pempheris vanicolensis
(not Cuvier, 1831):
Counts of holotype and non-types are given in Table
P. nyctereutes | P. sasakii | |||
---|---|---|---|---|
Holotype | Non-types | Holotype | Non-types | |
Number of individuals | 1 | 18 | 1 | 47 |
Standard length (mm) | 160.5 | 100.5–162.4 | 93.1 | 87.7–169.8 |
Dorsal fin rays | VI, 9 | VI, 9 | VI, 9 | VI–VII, 9–10 |
Anal fin rays | III, 44 | III, 42–44 | III, 43 | III, 40–46 |
Pectoral fin rays | 19 | 18–20 | 19 | 17–20 |
Left pored lateral-line scales | 79 | 72–81 | 72 | 67–78 |
Right pored lateral-line scales | 77 | 74–82 | 73 | 66–78 |
Scale above lateral line | 8 1/2 | 8 1/2–9 1/2 | 8 1/2 | 8 1/2–10 1/2 |
Scale rows below lateral line | 23 | 23–28 | 22 | 19–22 |
Circumpeduncular scales | 22 | 22–24 | damaged | 24 |
Gill rakers | 8+20 | 8+19–20 | 8+19 | 7–9+19–22 |
Widely distributed in the western Pacific Ocean excepting small oceanic islands and atolls in central and southeastern Pacific. In Japanese waters, this species is known from Yaizu in Shizuoka Prefecture, Iburi and Otsuki in Kochi Prefecture, Nagasaki in Nagasaki Prefecture, Uchinoura Bay in Kagoshima Prefecture, Hachijo-jima islands in Izu Islands, Tanega-shima to Yonaguni-jima islands in the Ryukyu Archipelago, Minamidaito-jima Island in the Daito Islands, Miyake-jima and Hachijo-jima islands in the Izu Islands, Haha-shima and Chichi-jima islands in the Ogasawara Islands. In Taiwanese waters, this species is known from Daxi in Yilan County, Yeh Liu in New Taipei City, Sihhu in Yunlin County, Tainan County, Checheng County, Hengchung, and Kenting in Pingtung County, Fugang in Taitung County, Lyudao, Lanyu, Xiao Liuqiu, and Penghu (Fig.
Although the taxonomic position of P. adusta was unsettled for a long time, the holotype (RMNH.PISC.6161: Ambon, Indonesia) matches well with the specimens in
The Maximum-Likelihood (ML) tree of Pempheris species recovered from mitochondria 16S and COI. Values of the Shimodaira–Hasegawa-like approximate likelihood ratio test (SH-aLRT) for the ML tree, bootstrap values (BS) of the NJ tree, and the posterior probability (PP) for the BI tree are indicated at the nodes unless the branch lengths are < 0.01. The museum voucher number of specimens are listed next to the taxon name.
Pempheris familia
Koeda & Motomura, 2017a: figs 1–3 (type locality: off Ototo-jima Island, Ogasawara Islands, Japan);
Pempheris japonica
(not Döderlein, 1883):
Counts of holotype and paratype are given in Table
Pempheris familia is a second species of the species group that is characterized by strongly ctenoid, adherent body scales with distinct basal and distal portions (see
Pempheris japonica
Döderlein, 1883: 125 (type locality: Tokyo, Japan);
Catalufa umbra Snyder, 1911: 528 (Misaki, Japan).
Catalufa japonica
(not Döderlein, 1883):
Pempheris umbra
(not Snyder, 1911):
Counts of syntypes and non-types are given in Table
Endemic to the region from southern Korea to southern Japan. In Japanese waters, P. japonica is distributed in the Pacific coast (north to Ishinomaki in Miyagi Prefecture, south to Kagoshima Prefecture), Japan Sea coast (east to Miyazu in Kyoto Prefecture, west to Tsuno-shima Island in Yamaguchi Prefecture), Tsushima Island, East China Sea coast (north to Nagasaki, south to Kagoshima prefectures), Miyake-jima and Hachijo-jima islands in Izu Islands, Tanega-shima, Yaku-shima, Iou-jima, Amami-oshima, and Okinawa-jima in Ryukyu Archipelago (very rare in the latter two islands) (Fig.
Pempheris nyctereutes
Jordan & Evermann, 1903: 339, fig. 14 (type locality: Taipei City [Hokoto], Taiwan);
Pempheris schwenkii
(not Bleeker, 1855):
Counts of holotype and non-types are given in Table
Recorded only from Taiwan, Hong Kong, and Vietnam. In Taiwanese waters, this species is known from Nang-fang-ao in Yilan County, Keelung City, Aodi, Wanli, and Gongliao in New Taipei City, Tainan County, Fugang in Taitung County, Hengchung and Maobitou in Pingtung County, Lyudao, and Penghu (Fig.
The taxonomic status of P. nyctereutes and P. sasakii that have similar morphology have been commonly confused. A comparison between these two species is discussed in the remarks for P. sasakii. Pempheris nyctereutes is sometimes collected by fisherman in southern Taiwan as bycatch with P. schwenkii and P. adusta, which are mainly distributed in coral-reef areas, suggesting that P. nyctereutes might be distributed in coral-reef areas. This species is also known from Ha Long Bay in Vietnam (FRLM 49700;
Pempheris oualensis
Cuvier in Cuvier & Valenciennes, 1831: 299 (type locality: Kosrae, Caroline Islands);
Pempheris otaitensis
(not Cuvier, 1831):
Pempheris
sp.:
Counts of holotype and non-types are given in Table
P. schwenkii | P. xanthoptera | |||
---|---|---|---|---|
Syntypes | Non-types | Holotype | Non-types | |
Number of individuals | 2 | 232 | 1 | 355 |
Number of individuals | 87.2, 89.7 | 20.4–125.9 | 116.8 | 28.2–136.6 |
Standard length | ||||
Dorsal fin rays | VI, 9 | VI–VII, 9–10 | VI, 9 | VI–VII, 9 |
Anal fin rays | III, 35–36 | III, 35–42 | III, 38 | III, 35–42 |
Pectoral fin rays | 17–18 | 16–18 | 18 | 16–19 |
Left pored lateral-line scales | 48 | 44–53 | 48 | 45–54 |
Right pored lateral-line scales | 48–49 | 45–53 | 47 | 46–54 |
Scale above lateral line | 3 1/2 | 3 1/2–4 1/2 | 3 1/2 | 3 1/2 |
Scale rows below lateral line | 10 | 10–13 | 10 | 10–13 |
Circumpeduncular scales | 12 | 10–12 | 12 | 12–14 |
Gill rakers | 7+18–19 | 6–9+18–22 | N/A | 7–9+18–21 |
Widely distributed in the western to central Pacific Ocean (not in the Hawaiian Islands), and Christmas Island and Cocos-Keeling Island in the Indian Ocean. In Japanese waters, this species is known from Tanega-shima to Yonaguni-jima islands in the Ryukyu Archipelago, Minamidaito-jima Island in the Daito Islands, and Haha-shima and Chichi-jima islands in the Ogasawara Islands. In Taiwanese waters, this species is known from Nang-fang-ao in Yilan County, Yeh Liu in New Taipei City, Tainan County, Chung-chou in Kaohsung County, Hengchung, and Kenting in Pingtung County, Fugang in Taitung County, and Lanyu (Fig.
This species has similar characters as Pempheris otaitensis Cuvier, 1831 and Pempheris ufuagari which share a distinct black blotch on pectoral-fin base and whose large body sizes reaches > 160 mm SL. However, P. oualensis is unique in having a dark coloration on the upper margin of its pectoral fin and a villiform tooth band extends outside the lips on large specimens. The scale count of this species varies among populations that specimens collected from the northwestern Pacific have 61–66 pored lateral-line scales, but specimens collected from southern Pacific have 67–71. Additionally, specimens collected from Andaman Sea have 5½ scale rows above lateral line, whereas those from the Pacific Ocean have 6½ or 7½, with the frequency of specimens with 7½ scale rows above lateral line being lower in the northwestern Pacific than in southern areas. In particular, the Andaman population may be a species distinct from the Pacific populations, but more specimens and genetic evidence are necessary to discuss whether the differences are interspecific or intraspecific.
Although
Liopempheris sasakii
Jordan & Hubbs, 1925: 228, pl. 10, fig. 1 (type locality: Toba, Mie Prefecture, Japan);
Pempheris sasakii:
Pempheris nyctereutes
(not Jordan & Evermann, 1903):
Counts of holotype and non-types are given in Table
Endemic to southern Japan known from Tateyama at Boso Peninsular in Chiba Prefecture, Misaki and Manazuru in Kanagawa Prefecture, Nishi-izu at Izu Peninsular in Shizuoka Prefecture, Toba and Shima in Mie Prefecture, Muroto, Susaki, Tosashimizu, Iburi, and Otsuki in Kochi Prefecture, Nobeoka and Nango in Miyazaki Prefecture, Uchinoura Bay, Kagoshima Bay, Ibusuki, and Minami-satsuma in Kagoshima Prefecture, Miyake-jima Island in Izu Islands, Tanega-shima, Yaku-shima, and Kuchinoerabu-jima islands in the northern Ryukyu Archipelago (Fig.
Pempheris sasakii has been commonly confused with P. nyctereutes and has been presumed to be the junior synonym (e.g.,
Pempheris sasakii is widely distributed in the Pacific coast of southern Japan, and commonly collected by set nets (but not abundant compared to P. xanthoptera). However, no specimens of this species have ever been collected from the Japan Sea coast, East China Sea coast, and the Ryukyu Archipelago. The distributions of both P. sasakii and P. nyctereutes are clearly isolated from each other (Fig.
Pempheris schwenkii
Bleeker, 1855: 314 (type locality: Batu Islands, Sumatera Utara Province, Indonesia);
Pempheris adusta
(not Bleeker, 1877):
Pempheris japonicus
(not Döderlein, 1883):
Pempheris oualensis
(not Cuvier, 1831):
Liopempheris vanicolensis
(not Cuvier, 1831):
Pempheris vanicolensis
(not Cuvier, 1831):
Pempheris xanthoptera
Tominaga, 1963 (in part paratypes): 287;
Pempheris
sp.:
Counts of holotype and paratypes are given in Table
Widely distributed in the western Pacific Ocean. In Japanese waters, this species is known from Tanega-shima to Yonaguni-jima islands in the Ryukyu Archipelago, Minamidaito-jima Island in Daito Islands, and very rarely collected from Minamisatsuma in Kagoshima Prefecture. In Taiwanese waters, this species is known from Daxi in Yilan County, Gungliau and Yeh Liu in New Taipei City, Chi-gu in Tainan County, Ke-tzu-liao in Kaohsiung County, Hengchung, and Dong-gang, Kenting in Pingtung County, Fugang in Taitung County, Lanyu, and Penghu (Fig.
Pempheris schwenkii has been thought to be widely distributed in the Indo-Pacific Ocean. However, our genetic study revealed the interspecific difference between specimens from the Indian and Pacific oceans, and southern Japan.
Similarly, the Pacific species and the southern Japanese species also showed significant difference between specimens from south of the Ryukyu Archipelago and specimens from mainland Japan (unpublished data). These two are clearly different species, because both species are distributed in the Osumi Islands (Tanega-shima, Yaku-shima, and Kuchinoerabu-jima islands; Fig.
On the basis of the taxonomic confusion between P. schwenkii and P. xanthoptera, the standard Japanese name “Minami-hatampo” was used for both species, and recently, P. schwenkii was tentatively recognized as having “Pacific” and “southern Japan” types, the latter closely matching P. xanthoptera sensu
The juveniles of P. schwenkii were collected from Minami-daito Island in the Daito Islands. Pempheris ufuagari is known as an endemic species which is found in the Daito and Ogasawara islands (see below), meaning that interaction between the species can occur at these localities. However, P. schwenkii and P. xanthoptera have never been collected from the Ogasawara and Daito islands, respectively, indicating that the P. ufuagari and P. schwenkii group (with P. xanthoptera) may have a different dispersal strategy.
Pempheris ufuagari
Pempheris oualensis
(not Cuvier, 1831):
Pempheris otaitensis
(not Cuvier, 1831):
Counts of holotype and paratypes are given in Table
Endemic to the Daito and Ogasawara islands (Fig.
Pempheris ufuagari is most similar to P. otaitensis, known only from French Polynesia and Samoa, sharing a distinct black blotch on the pectoral-fin base, yellow dorsal and caudal fins, and a blackish band on the anal-fin outer margin. However, the former can clearly be distinguished from the latter in having 62–71 pored lateral-line scales (vs 69–79 in P. otaitensis), 6½–7½ scale rows above lateral line (vs 8½), 37–43 predorsal scales (vs 44–48), and the tip of the dorsal fin blackish (vs anterior margin to tip blackish).
Pempheris vanicolensis
Cuvier, 1831: 305 (type locality: Vanikoro Island, Santa Cruz Islands);
Pempheris
sp.:
Counts of holotype and non-types are given in Table
Widely distributed in the western Pacific Ocean except for small islands and atolls in central Pacific. In Japanese waters, this species is known from Iriomote-jima and Yonaguni-jima islands in the southern Ryukyu Archipelago. In Taiwanese waters, this species is known from Yeh Liu and Wang-li in New Taipei City, and Hengchung and Kenting in Pingtung County (Fig.
This species was described by
Pempheris xanthoptera
Tominaga, 1963: 286, fig. 12 (type locality: Manazuri, Kanagawa Prefecture, Japan);
Pempheris molucca (not Cuvier, 1829): Temminck and Schlegel 1844: 85, pl. 44, fig. 3.
Pempheris oualensis
(not Cuvier, 1831):
Pempheris schwenkii
(not Bleeker, 1855):
Pempheris vanicolensis
(not Cuvier, 1831):
Pempheris japonica (not Döderlein, 1883): Nakamura 1993: 148, fig. 6.
Counts of of holotype and paratypes are given in Table
Endemic to the Northwest Pacific, recorded only from Japan, Jeju Island in Korea (
Although
The standard Japanese name “Minami-hatampo” was used for both species and caused confusion, but this name should be adopted for P. schwenkii (see remarks of P. schwenkii). Therefore, a new standard Japanese “Mizuho-hatampo” is proposed for P. xanthoptera. “Mizuho” is an alternative name for Japan that frequently appears in ancient Japanese mythology and poetry, and it derives from the fact that the species is primarily distributed across the Japanese mainland.
We are especially grateful to K. Tachihara, T. Yoshino, H. Imai (University of the Ryukyus), H. Motomura (The Kagoshima University Museum), and H.-C. Ho (National Kaohsiung University of Science and Technology; previously National Museum of Marine Biology & Aquarium) for their supervision and many useful suggestions for this research. We are greatly appreciative of the assistance of the following individuals and institutions for allowing us to examine types and non-type specimens, and kind support during visits of KK and loans of specimens: K.-T. Shao and P.-L. Lin (ASIZP), O. Crimmen and P. Campbell (NHMUK), H. Endo and M. Sato (BSKU), J.E. Randall, A. Suzumoto, and L. O’Hara (BPBM), H. Kawase (CBM), M. Miya, T. Sado, and H. Kobayashi (CMNH), T. Nakabo, A. Tokairin, and N. Nakayama (FAKU), S. Kimura and Y. Hibino (FRLM), T. Mezaki and T. Toshino (KBF), H. Senou (KPM), P. Pruvost, G. Duhamel, R. Causse, and G. Zora (MNHN), K. Matsuura, G. Shinohara, M. Nakae (NSMT), S. Oka, K. Miyamoto, and N. Hanahara (OCF), R. Ruiter and M.J.P. van Oijen (RMNH), J. Williams and S. Raredon (USNM), K. Hayashi and K. Hagiwara (YCM), P. Bartsch and C. Lamour (ZMB), R. Thiel and I. Eidus (ZMH), and M. Aizawa, K. Sakamoto, and R. Ueshima (ZUMT). We also thank members, collaborators, and specimen providers of Tachihara Laboratory and Koeda Laboratory at University of the Ryukyus, KAUM, National Museum of Marine Biology & Aquarium, Kuroshio Biological Research Foundation, and ZUMT. H. Senou provided data of types at NMW. G. S. Hardy (Ngunguru, New Zealand) kindly improved the English in the manuscript.
The authors have declared that no competing interests exist.
No ethical statement was reported.
This study was supported in part by JSPS KAKENHI 21K06313 JP and 21K15144 JP, the Sasakawa Scientific Research Grant from The Japan Science Society, JST CREST (JPMJCR23J2) and FOREST Program (JPMJFR214D), Ocean Shot from the Sasakawa Peace Foundation, a Grant-in-Aid from the Japan Society for the Promotion of Science for JSPS Fellows (DC1: 23-2553; PD: 21J01755), and JSPS Overseas Research Fellowships (29-304).
Conceptualization: KK. Data curation: MBU, KK. Formal analysis: MBU, KK. Funding acquisition: KK, MBU. Project administration: KK. Validation: MBU. Visualization: KK. Writing - original draft: KK. Writing - review and editing: KK, MBU.
Keita Koeda https://orcid.org/0000-0003-3932-3002
Manabu Bessho-Uehara https://orcid.org/0000-0002-7388-464X
All of the data that support the findings of this study are available in the main text or Supplementary Information.
Preserved specimens of genus Pempheris examined in the present study
Data type: xlsx
requency distributions of pored lateral-line scales of species of Pempheris distributed in Japan and Taiwan, with P. flavicycla for comparison
Data type: xlsx