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Research Article
A review of the genus Pempheris (Teleostei, Pempheridae) found in Japan and Taiwan
expand article infoKeita Koeda, Manabu Bessho-Uehara§
‡ University of the Ryukyus, Okinawa, Japan
§ Tohoku University, Sendai, Japan
Open Access

Abstract

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.

Key words

Distribution, morphology, Pempheris sasakii, Pempheris xanthoptera, sweepers, taxonomy

Introduction

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 (Tominaga 1963). The genus Pempheris was first proposed by Cuvier (1829) for Pempheris touea Cuvier, 1829, which is presently recognized as a junior synonym of Pempheris compressa (Shaw, 1790) (Tominaga 1968). In total, 84 nominal species have been described for the genus to date (Fricke et al. 2024), with almost half of them newly described from the western Indian Ocean from 2014 to 2015 (e.g., Randall and Victor 2014; Randall and Victor 2015). Because the validity of these Indian Ocean species needs to be reassessed, the taxonomy of the whole genus has long been confused due to difficulties in identifying its species.

The taxonomy of the family Pempheridae in Japanese waters was reviewed by Tominaga (1963) based on a comparison of morphology, and he recognized four species of the genus Pempheris from Japan. Since then, two new species have been described: Pempheris familia Koeda & Motomura, 2017 and Pempheris ufuagari Koeda, Yoshino & Tachihara, 2013, and two new-to-Japan species were reported, Pempheris oualensis Cuvier, 1831 and Pempheris vanicolensis Cuvier, 1831, from the Japanese waters (Koeda et al. 2010a, b, 2013a; Koeda and Motomura 2017a). However, the validity of Pempheris sasakii Jordan & Hubbs, 1925 and Pempheris xanthoptera Tominaga, 1963, both described from Japan, have never been evaluated. Therefore, the present study conducted a direct comparison of the morphology, including of the type specimens of Pempheris nyctereutes Jordan & Evermann, 1902 and Pempheris schwenkii Bleeker, 1877. Although many species of the genus commonly distributed in Japan are also found in Taiwan, the taxonomy of the genus in Taiwan has never been reviewed, and misidentifications are occasionally observed in the Taiwanese literature. Here, we reviewed the species of the genus Pempheris that occurred in Japan and Taiwan with detailed descriptions, based on the type specimens and large numbers of non-types, providing diagnoses, identification keys, and distributional ranges. Additionally, the published literature which relates to the genus Pempheris of Japan and Taiwan was re-examined and corrected as much as possible.

Materials and methods

Preserved materials examined in the present study including large numbers of specimens collected by KK are listed in Suppl. material 1. Comparative materials of the type specimens of species of the genus Pempheris are listed in Koeda et al. (2013a, 2014) and Koeda and Motomura (2017a). Counts and measurements followed Koeda et al. (2014). All measurements were made on the left side when possible, using digital calipers and rounded to the nearest 0.1 mm. Standard and head lengths are abbreviated as SL and HL, respectively. Osteological characters, including vertebral counts, were observed from radiographs. The descriptions of general morphology shared by species of the genus are not repeated here. Data of type specimens are given in parentheses. The distribution maps were made based on the specimens examined and collected by KK, quality underwater photographs, and literature records with a good illustration and/or sufficient diagnostic information to provide for positive identification. Photographs were taken by KK except when a credit line is given. Synonym lists for each species are shown only for the related references for Japan and Taiwan and the original descriptions. Institutional codes used in this study follow Fricke et al. (2024) with an addition: University of the Ryukyus, Ichthyological Laboratory (URIL).

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 Koeda et al. (2014). We concatenated and aligned the obtained 16S and COI sequences using MAFFT alignment (v. 7.490) (Katoh and Standley 2013) with default settings in Geneious Prime software (v. 2023.2.1) (Biomatters). Uncorrected pairwise distances (p-distances) among the sequences of different specimens were estimated using Geneious Prime software. Phylogenetic relationships were inferred using maximum-likelihood (ML) inference, neighbor-joining (NJ), and Bayesian inference (BI) methods. The ML tree was reconstructed using IQ-TREE 1.6.12 (Trifinopoulos et al. 2016) with a partitioned model for 16S (sites 1–481) and COI (sites 482–837). The best-fit substitution model was chosen by ModelFinder with “Auto” option which automatically selects the best-fit model for each partition. We performed the Shimodaira–Hasegawa-like approximate likelihood ratio test (SH-aLRT) 1,000 times to assess the nodal support (Guindon et al. 2010). The NJ tree was reconstructed using the Tamura-Nei model, and a consensus tree was generated by resampling 10,000 replicates of bootstrap analysis. The BI tree was constructed with MrBayes 3.2.6 (Huelsenbeck and Ronquist 2001), using the GTR substitution model and the invgamma rate variation model. Four independent Markov chain Monte Carlo (MCMC) runs were conducted for 2,100,000 generations, subsampling trees every 200 cycles, with the initial 100,000 trees discarded as burn-in. The majority rule consensus of the remaining trees was used to determine clade posterior probabilities.

Taxonomic account

Pempheris Cuvier, 1829

Standard Japanese name: Hatampo-zoku

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).

Description

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.

Distribution

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 (Mouneimne 1979; Golani and Ben-Tuvia 1986; Golani and Diamant 1991; Koeda et al. 2014).

Remarks

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.

Fowler (1906), Snyder (1911), and Ogilby (1913) attempted to divide the genus Pempheris into two genera based on scale morphology. Tominaga (1968) described and compared the internal anatomy of many species of the genus and suggested that several species of genus Pempheris possess transitional characteristics to the genus Parapriacanthus. Therefore, he subdivided the genus Pempheris into seven groups. That work indicated that a systematic revision of the genus should be pursued. Although the authors of the present study are now revising the systematic taxonomy of the family Pempheridae on the basis of morphology and molecular approaches, the present classification of the genus Pempheris is tentatively used only for the species of Japan and Taiwan.

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.

Key to the species of genus Pempheris in Japan and Taiwan (with distributions in parentheses after the species name)

1 12–15 scale rows above lateral line; scales on lateral body strongly ctenoid and adherent, with distinct basal and distal portions (Koeda et al. 2013a: fig. 2b); ventral surface of abdomen rounded, cross-sectional outline U-shaped; coracoid slightly expanded posteriorly; large ventral fenestra between coracoid and cleithrum 2
3½–10½ scale rows above lateral line; scales on lateral body weakly ctenoid and deciduous (Koeda et al. 2013a: fig. 2a), semicircular in shape; ventral surface of abdomen slightly or well keeled, cross-sectional outline V-shaped; coracoid enormously expanded posteriorly; very small ventral fenestra between coracoid and cleithrum 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

Figs 1, 2, Suppl. material 2 Standard Japanese name: Ryukyu-hatampo

Pempheris adusta Bleeker, 1877: 50, pl. 383, fig. 1 (type locality: Ambon Island, Molucca Islands, Indonesia); Koeda et al. 2013a: 235; Koeda et al. 2013b: 221, fig. 1; Koeda et al. 2013c: 123, fig. 1; Koeda et al. 2014: 303, fig. 1; Motomura and Matsuura 2014: 270, unnumbered figs; Koeda and Motomura 2015: 139, fig. 1; Koeda et al. 2015: 279; Kaburagi 2016: 98, upper fig. (without scientific name; shown as “Ryukyu-hatampo” in Japanese); Koeda et al. 2016a: 519; Koeda et al. 2016b: 50, fig. 224; Koeda et al. 2016c: 8, fig. 3-G; Koeda and Motomura 2017a; Koeda 2017a: 5, fig. 1 (middle fig.); Kimura et al. 2017: 119, fig. 5; Planning and Tourism Division of Kikai Town 2017: 4, unnumbered figs; Nakae et al. 2018: 266; Koeda 2018a: 193, unnumbered figs; Koeda 2018b: 298, unnumbered fig.; Koeda 2018c: 340, unnumbered figs; Mochida and Motomura 2018: 30; Koeda 2019: 926, unnumbered figs; Murase et al. 2019: 132, fig. 283; Fujiwara and Motomura 2020: 28; Koeda 2020a: 407, unnumbered figs; Koeda 2020b: 926, unnumbered figs; Motomura and Uehara 2020: 45; Murase et al. 2021: 166, fig. 339; Koeda et al. 2022: 5; Motomura 2023: 126.

Pempheris mangula (not Cuvier, 1829): Schmidt 1913: 121; Randall and Lim 2000: 622.

Pempheris oualensis (not Cuvier, 1831): Snyder 1912: 497; Okada 1938: 179; Okada and Matsubara 1938: 179; Aoyagi 1948: 49; Matsubara 1955: 590; Tominaga 1963: 289; Honda 1972: 72; Masuda et al. 1975 (in part): 199, pl. 33-D; Yoshino et al. 1975: 75; Hayashi 1984 (in part): 160, pl. 151-E; Shen 1984: 74, pl. 74, fig. 334-1; Shao et al. 1992: 177, unnumbered fig.; Shen 1993: 390, pl. 114 (fig. 1); Shao and Chen 1991: 162, unnumbered fig.; Chen et al. 1995 (probably in part): 25; Mochizuki 1995: 389; Yoshigou et al. 2001: 141; Chen 2003 (in part): 134; Ito 2009: 80, unnumbered fig.; Shao et al. 2008: 254; Chen et al. 2010: 265, fig. D; Chang et al 2011: 46; Shen and Wu 2011: 498, unnumbered fig; Shao et al. 2013 (in part): 163, unnumbered fig. (upper left); Chiang et al. 2014: 183, unnumbered fig.

Pempheris sp.: Uchida 1933: 218 (in part); Senou et al. 2006a: 77; Senou et al. 2007: 56; Hatooka 2002 (in part): 878; Senou et al. 2002: 212; Yoshino 2008: 211, unnumbered figs (lower two); Koeda et al. 2010a: 75; Motomura et al. 2010 (in part): 131; Koeda et al. 2012a: 71; Koeda et al. 2012b: 1086; Hatooka and Yagishita 2013 (in part): 984; Motomura et al. 2013: 169, unnumbered figs.

Pempheris vanicolensis (not Cuvier, 1831): Chen et al. (2010): 266, fig. B.; Shen and Wu 2011: 498, unnumbered fig.

Diagnosis

Counts of holotype and non-types are given in Table 1 of Koeda et al. (2013b). Dorsal-fin spines 5 or 6, very rarely 5, soft rays 8–10, very rarely 8 or 10; anal-fin spines 3, soft rays 37–45, usually > 40; pectoral-fin rays 16–19, usually 17 or 18; pored lateral-line scales 51–62, usually > 54; scale rows above lateral line 4½–5½ (usually 4½); scale rows below lateral line 11–16, usually 12–14; predorsal scales 26–38; circumpeduncular scales 12–18, usually 16; gill rakers 7–10+20–23 = 28–32, usually 8–9+20–22 = 29–31; head length 26.3–31.8% SL; body depth 40.2–47.3% SL; eye diameter 36.0–47.1% HL; upper jaw length 48.1–57.1% HL; maximum 182.7 mm SL, usually < 160 mm SL; scales weakly ctenoid, deciduous, thin, semicircular in shape, far wider than long; body copper to brownish, whiter in nighttime; faint blackish blotch on pectoral-fin base; tip and/or anterior margin of dorsal fin blackish; blackish band on outer margin of anal fin usually absent; blackish or dusky band on posterior edge of caudal fin; narrow band of villiform teeth in jaws; abdomen cross-sectional outline V-shaped.

Table 1.

Counts of Pempheris nyctereutes and P. sasakii.

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

Distribution

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. 2). Common in coral-reef areas of the Ryukyu Archipelago of Japan, the southern coast, and eastern islands (Lyudao and Lanyu) of Taiwan, but few in other areas. Specimens collected from 0–20 m depth.

Figure 1. 

Pempheris adusta A fresh specimen (ZUMT 62301, Nishidomari, Otsuki, Kochi, Japan) and B underwater photograph (Maeda, Onna, Okinawa-jima Island, Japan).

Figure 2. 

Distribution of Pempheris adusta based on the collection localities of specimens.

Remarks

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 Koeda et al. (2013b). The original description of the species is also supported as follows: figures of six species (P. mangula Cuvier, 1829, P. schwenkii, P. vanicolensis, P. adusta, P. otaitensis Cuvier, 1831, and P. oualensis) were illustrated in the plate of Bleeker (1877); the first three species have no black blotch and the latter three species possess a black blotch on the pectoral-fin base. This was also mentioned in the text descriptions, where he specified that P. oualensis and P. otaitensis had a black blotch, but P. adusta has a black or brown blotch on the pectoral-fin base visible in the figure and clearly corresponding with the descriptions of the three species. Koeda et al. (2014) indicated that Pempheris adusta was widely distributed species from the Indian to Pacific oceans and had intraspecific variations in their morphology. The Pacific group differs from the Indian Ocean group in the following characters and individuals from Andaman Sea have characters intermediate between these two groups: pored lateral-line scales 51–62 (vs 56–63 in Indian Ocean; 53–57 in Andaman Sea); scale rows above lateral line usually 4½ (vs usually 5½ in Indian Ocean; 4½ in Andaman Sea); usually no blackish band on anal fin (vs distinct blackish band on outer margin of anal fin in Indian Ocean and Andaman Sea); blackish band on anal-fin base (no band on anal-fin base in Indian Ocean and Andaman Sea); and anterior margin of dorsal fin blackish (tip of dorsal fin blackish in Indian Ocean and Andaman Sea). Furthermore, Koeda et al. (2014) demonstrated that nucleotide sequences of specimens collected from the Red Sea, the Andaman Sea, and the Pacific Ocean showed only 0.4% difference in mitochondrial 16S ribosomal DNA. Coupled with the observation of small morphological differences, they considered Pempheris flavicycla Randall, Satapoomin & Alpermann, 2014 (type locality: Mafia Island, Chole Islands, Chole Bay, Tanzania) to be a junior synonym of P. adusta. However, Randall et al. (2014) countered this opinion based on the 2.5% difference in COI sequences, stating that P. adusta is a species in Pacific Ocean, P. flavicycla is the valid species in Indian Ocean, and described the Andaman group as a subspecies Pempheris flavicycla marisrubri Randall, Bogorodsky & Alpermann, 2014. Based on the genetic comparison incorporating the COI and 16S genes in the present study, it was shown that there is more than a 2% genetic difference between P. adusta and P. flavicycla (Fig. 3), which is not a subtle difference when compared to differences among other similar species in the genus. Although the issue of overlapped morphological differences remains in P. adusta and P. flavicycla, it is reasonable to support the opinion of Randall et al. (2014) that P. flavicycla is considered as a valid species distributed in Indian Ocean at the present time.

Figure 3. 

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, 2017

Figs 4, 5, Suppl. material 2 Standard Japanese name: Bonin-hatampo

Pempheris familia Koeda & Motomura, 2017a: figs 1–3 (type locality: off Ototo-jima Island, Ogasawara Islands, Japan); Koeda 2018b: 299, unnumbered fig.

Pempheris japonica (not Döderlein, 1883): Toyama 1937: 36; Kurata et al. 1971: 25; Zama and Fujita 1977: 102; Randall et al. 1997: 35; Hatooka and Yagishita 2013: 983 (in part); Koeda 2017a: 6, fig. 1 (lower fig.).

Diagnosis

Counts of holotype and paratype are given in Table 1 of Koeda and Motomura (2017a). Dorsal-fin rays VI, 9–10; anal-fin rays III, 35–36; pectoral-fin rays 17; pored lateral-line scales 84–88; scale rows above lateral line 14–15; scale rows below lateral line 28–30; predorsal scales 50–55; circumpeduncular scales 26; gill rakers 12–13+22–26 = 34–39; head length 28.8–30.1% SL; body depth 42.9–43.7% SL; eye diameter 46.8–47.6% HL; upper jaw length 50.0–56.3% HL; maximum 153 mm SL; scales strongly ctenoid, adherent, divided into basal and distal halves (Koeda et al. 2013a: fig. 2b); body copper; distinct blackish blotch on pectoral-fin base; tip of dorsal and anal fins broadly black, and rest brown; narrow band of villiform teeth in jaws; abdomen cross-sectional outline U-shaped.

Figure 4. 

Pempheris familia A fresh specimen (KAUM–I. 74713, 153.1 mm SL, holotype, Ototo-jima Island, Ogasawara Islands, Japan, photo taken by K. Kuriiwa) and B underwater photograph (Ani-jima Island, Ogasawara Islands, Japan).

Distribution

Endemic to the Ogasawara Islands (Fig. 5).

Figure 5. 

Distribution of Pempheris familia (diamond) and Pempheris japonica (triangles and star for type locality) based on the collection locality of the specimens.

Remarks

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 Koeda et al. 2013a: fig. 2b) in the Northern Hemisphere shared only with P. japonica (Koeda et al. 2012a, 2013a; Koeda and Motomura 2017a). The remaining six species of this group are endemic to Australia, New Zealand, or French Polynesia (Mooi and Jubb 1996; Mooi 1998, 2000). Although P. familia shares most morphological characteristics with P. japonica, the former can be clearly distinguished from the latter in scale counts and the distinct black blotch on pectoral-fin base (Koeda and Motomura 2017a).

Pempheris japonica Döderlein, 1883

Figs 5, 6, Suppl. material 2 Standard Japanese name: Tsumaguro-hatampo

Pempheris japonica Döderlein, 1883: 125 (type locality: Tokyo, Japan); Jordan et al. 1913: 137; Tanaka 1931: 25; Uchida 1933: 208; Okada 1938: 179; Okada and Matsubara 1938: 179; Matsubara 1955: 590; Tominaga 1963: 278, fig. 6; Takemura and Yasuda 1965: 159; Masuda et al. 1975: 199, pl. 33-B; Hayashi 1984 (in part): 160, pl. 151-C; Kohno 1986: 135, fig. 1; Masuda and Kobayashi 1994: 180, fig. 4; Mochizuki 1995: 389, unnumbered fig.; Hatooka 1997: 380, unnumbered fig.; Hatooka 2002 (in part): 877; Senou et al. 2002: 212; Senou et al. 2006b: 463; Aramata 2007: 171, unnumbered fig.; Yoshino 2008: 211, unnumbered fig.; Koeda et al. 2010a: 74; Koeda et al. 2010b: 81; Motomura et al. 2010: 131; Takagi et al. 2010: 69, unnumbered figs; Kohno et al. 2011: 208, unnumbered fig.; Koeda et al. 2012a: 65; Hatooka and Yagishita 2013 (in part): 983; Koeda et al. 2013b: 235; Motomura et al. 2013: 168, unnumbered fig.; Kawano et al. 2014: 48; Koeda et al. 2014: 327; Koeda and Motomura 2015: 139; Koeda et al. 2015: 275; Ikeda and Nakabo 2015: 159, figs 5–7; Takeuchi et al. 2015: 8; Koeda 2017b: 190, unnumbered fig.; Koeda and Motomura 2017a; Fujiwara et al. 2018: 58, Fig. 8L; Kagoshima City Aquarium Foundation 2018: 210, unnumbered fig.; Koeda 2018b: 298, unnumbered fig.; Nakae et al. 2018: 266; Koeda 2018c: 341, unnumbered figs; Murase et al. 2019: 132, fig. 282; Koeda 2020a: 408, unnumbered figs; Sonoyama et al. 2020: 78; Murase et al. 2021: 166, fig. 338; Koeda 2022: 158, unnumbered fig.; Koeda et al. 2022: 6; Motomura 2023: 127; Sakurai et al. 2024: 77, fig. 4D.

Catalufa umbra Snyder, 1911: 528 (Misaki, Japan).

Catalufa japonica (not Döderlein, 1883): Jordan and Hubbs 1925: 227.

Pempheris umbra (not Snyder, 1911): Okada 1938: 179; Okada and Matsubara 1938: 179; Matsubara 1955: 59, pl. 54 (fig. 189); Takemura and Yasuda 1965: 159; Abekawa and Nishi 1969: 24.

Diagnosis

Counts of syntypes and non-types are given in Table 1 of Koeda and Motomura (2017a). Dorsal-fin rays VI, 10–12; anal-fin rays III, 34–40; pectoral-fin rays 16–17; pored lateral-line scales 69–82; scale rows above lateral line 12–13; scale rows below lateral line 26–30; predorsal scales 40–44; circumpeduncular scales 22–24; gill rakers 8–12+19–25 = 28–35; head length 28.3–31.4% SL; body depth 43.2–47.7% SL; eye diameter 38.1–50.0% HL; upper jaw length 50.0–56.3% HL; maximum 153 mm SL; scales strongly ctenoid, adherent, divided into basal and distal halves (see Koeda et al. 2013a: fig. 2b); body copper; no or faint blackish blotch on pectoral-fin base; tip of dorsal and anal fins broadly black, remainder brown; narrow band of villiform teeth in jaws; abdomen cross-sectional outline U-shaped.

Distribution

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. 5).

Remarks

Döderlein (1883) described P. japonica based on syntypes collected from Tokyo Bay, and Snyder (1911) described Catalufa umbra based on the holotype collected from Kanagawa Prefecture. The type specimens of the two nominal species were compared in the present study, and no differences were observed. Therefore, C. umbra is confirmed as a junior synonym of P. japonica, in agreement with Tominaga (1963).

Snyder (1912) and Shao et al. (2008) reported P. japonica in the fish checklists of Okinawa Island and southern Taiwan, respectively. However, the specimens they observed and identified as P. japonica (CAS-SU 22002; ASIZP 61383) are identified as P. schwenkii and P. vanicolensis, respectively (Koeda et al. 2012a). Subsequently, Randall et al. (1997) reported P. japonica from the Ogasawara Islands. However, that report was not based on specimens and/or underwater observations (refer to pers. comm. of R. Mooi on p. 35) and was probably the misidentification of P. familia which is similar to P. japonica. Although Hayashi (1984) and Hatooka (2000, 2002) included the Philippines, and Hatooka and Yagishita (2013) included Taiwan in the distributional range of P. japonica, specimens of P. japonica from these localities have never been discovered. Our results indicate that P. japonica is not distributed in these localities, and the species is endemic to Japan and southern Korea. Compared to the distribution of other species of the same genus found in the Northern Hemisphere, this species can be said to be the most temperate species adapted to the lowest water temperatures. Until the 2010s, this species was distributed only as far as the Boso Peninsula, a trend known for many tropical fish species. However, in recent years, new distribution records have been reported from Fukushima and Miyagi Prefectures, likely a result of northward range expansion due to global warming.

Figure 6. 

Fresh specimen of Pempheris japonica, KAUM–I. 89834, 125.0 mm SL, Nakakoshiki-jima Island, Koshiki Islands, Japan.

Pempheris nyctereutes Jordan & Evermann, 1903

Figs 7, 8; Table 1, Suppl. material 2 Standard Japanese name: Taiwan-hatampo

Pempheris nyctereutes Jordan & Evermann, 1903: 339, fig. 14 (type locality: Taipei City [Hokoto], Taiwan); Okada 1938: 179; Okada and Matsubara 1938: 179; Matsubara 1955:590; The Marine Ecological Researching Society of Kagoshima University 1966: 19; Shen 1993: 390, unnumbered fig.; Koeda et al. 2013a: 237; Koeda et al. 2014: 327; Tominaga 1963: 281, fig. 8; Hayashi 1984 (in part): 161; Randall and Lim 2000: 622; Hatooka 2002 (in part): 877; Hatooka and Yagishita 2013 (in part): 984; Shen and Wu 2011: 497, unnumbered fig.; Chiang et al. 2014: 183, unnumbered fig.; Koeda 2019: 927, unnumbered figs; Koeda 2020b: 927, unnumbered figs; Koeda et al. 2022: 9.

Pempheris schwenkii (not Bleeker, 1855): Lee 1996: 97, unnumbered fig.

Diagnosis

Counts of holotype and non-types are given in Table 1. Dorsal-fin rays VI, 9; anal-fin rays III, 42–44; pectoral-fin rays 18–20; pored lateral-line scales 72–81 usually > 79; scale rows above lateral line 8 1/2–9 1/2; scale rows below lateral line 23–28; circumpeduncular scales 22–24; gill rakers 8+19–20 = 27–28; head length 30.0–30.9%; body depth 44.6–46.9%; eye diameter 41.0–46.4%; upper jaw length 51.3–53.5%; maximum 161 mm SL; snout sharp; scales weakly ctenoid, deciduous, thin, semicircular in shape, far wider than long (see Koeda et al. 2013a: fig. 2a); body silver to dark brown in fresh specimens; tip and anterior margin of dorsal fin zonally blackish; anal fin pale with faint black band on base; paired fins pink; posterior half of caudal fin dusky; body light brown in fixed specimen; black pigmentation on each fin usually persistent, but not in holotype; no blackish blotch on pectoral-fin base; narrow band of villiform teeth in jaws; abdomen cross-sectional outline is V-shaped.

Distribution

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. 8).

Figure 7. 

Fresh specimen of Pempheris nyctereutes, NMMB-P 27469, 149.4 mm SL, Penghu, Taiwan.

Figure 8. 

Distribution of Pempheris nyctereutes (solid triangles and solid star for holotype locality) and Pempheris sasakii (solid diamonds and open star for holotype locality) based on the collection localities of the specimens.

Remarks

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; Koeda 2018d) and Hong Kong (BMNH 1939.3.23.48). Okada (1938), Okada and Matsubara (1938), and Matsubara (1955) admitted P. nyctereutes and P. sasakii as valid species being endemic to Taiwan and Japan, respectively, and Okada (1938) gave the Japanese name “Taiwan-hatampo” and “Mie-hatampo” to them. Tominaga (1963) later proposed “Takasago-hatampo” for P. nyctereutes as a new Japanese name without any reasons (probably overlooking the original designation), and Koeda et al. (2022) followed that. Based on Rule 6 of the guidelines for the naming of standard Japanese names for fishes (The Ichthyological Society of Japan 2021), “Taiwan-hatampo” proposed by Okada (1938) should be adopted for P. nyctereutes.

Pempheris oualensis Cuvier, 1831

Figs 9, 10, Suppl. material 2 Standard Japanese name: Yume-hatampo

Pempheris oualensis Cuvier in Cuvier & Valenciennes, 1831: 299 (type locality: Kosrae, Caroline Islands); Masuda et al 1975 (in part): 199; Hayashi 1984 (in part): 161; Chen et al. 1995 (probably in part): 25; Shao et al. 2008: 254; Koeda et al. 2010a: 72, fig. 1; Koeda et al. 2010b: 81; Koeda et al. 2012a: 71; Koeda et al. 2013a: 231, fig. 1b; Koeda et al. 2013b: 126; Koeda et al. 2014: 327; Koeda and Motomura 2015: 139; Koeda et al. 2015: 276, fig. 1; Koeda et al. 2016b: 50, fig. 225; Koeda and Motomura 2017a; Koeda 2017a: 7, fig. 2 (upper fig.); Kimura et al. 2017: 119, fig. 7; Planning and Tourism Division of Kikai Town 2017: 4, unnumbered figs; Nakae et al. 2018: 266; Koeda 2018a: 194, unnumbered figs; Koeda 2018b: 298, unnumbered fig.; Mochida and Motomura 2018: 30; Koeda 2019: 928, unnumbered figs; Motomura and Uehara 2020: 45; Koeda 2020b: 928, unnumbered figs; Motomura 2023: 129.

Pempheris otaitensis (not Cuvier, 1831): Schmidt 1913: 121; Randall et al. 1997 (in part): 35, pl. 8 (fig. F).

Pempheris sp.: Hatooka 2002 (in part), 878; Motomura et al. 2010 (in part): 131, fig. 253; Hatooka and Yagishita 2013 (in part): 984.

Diagnosis

Counts of holotype and non-types are given in Table 2 of Koeda et al. (2010a). Dorsal-fin rays VI, 9–10; anal-fin rays III, 38–46; pectoral-fin rays 17–19; pored lateral-line scales 60–72; scale rows above lateral line 5 1/2–7 1/2, usually 6 1/2; scale rows below lateral line 13–17; predorsal scales 33–44; circumpeduncular scales 18–22; gill rakers 8–9+20–22 = 28–31; head length 27.7–31.5%; body depth 40.9–48.2%; eye diameter 36.4–43.9%; upper jaw length 50.0–56.0%; maximum 208 mm SL, usually < 180 mm SL; scales weakly ctenoid, deciduous, thin, semicircular in shape, far wider than long (Koeda et al. 2013a: fig. 2a); body silver; distinct blackish blotch on pectoral-fin base; anterior margin of dorsal fin zonally blackish; blackish band on anal-fin base with rarely blackish band on its margin; upper margin of pectoral fin dusky; villiform tooth band extending outside lips on large specimen; abdomen cross-sectional outline V-shaped.

Table 2.

Counts of Pempheris schwenkii and P. xanthoptera.

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

Distribution

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. 10).

Figure 9. 

Pempheris oualensis A fresh specimen (NMMB-P 27821, 175.4 mm SL, Hengchung, Pingtung, Taiwan) and B underwater photograph (Dobuiso, Ogasawara Islands, Japan).

Figure 10. 

Distribution of Pempheris oualensis based on the collection locality of the specimens.

Remarks

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 Okada (1938) gave the Japanese name “Ryukyu-hatampo” to P. oualensis which was listed in Snyder (1912), specimen (USNM 75468) used for the list was re-identified to P. adusta (Koeda et al. 2013b). Therefore, the standard Japanese name “Ryukyu-hatampo” was adopted for P. adusta and the standard Japanese name “Yume-hatampo” was provided to P. oualensis by Koeda et al. (2010a) who first reported this species from Japanese waters. This species is the largest species of Pempheris that reaches > 200 mm SL; the largest specimen was collected from the Ogasawara Islands and measured 208.8 mm SL (KAUM–I. 74584).

Pempheris sasakii Jordan & Hubbs, 1925

Figs 8, 11, Suppl. material 2 Standard Japanese name: Mie-hatampo

Liopempheris sasakii Jordan & Hubbs, 1925: 228, pl. 10, fig. 1 (type locality: Toba, Mie Prefecture, Japan); Tanaka 1931: 25.

Pempheris sasakii: Uchida 1933: 217; Okada 1938: 179; Okada and Matsubara 1938: 180; Matsubara 1955: 590; Tominaga 1963: 283, fig. 10; Koeda et al. 2013a: 231; Koeda et al. 2014: 327.

Pempheris nyctereutes (not Jordan & Evermann, 1903): Hayashi 1984 (in part): 160, pl. 350-G; Hatooka 2002 (in part): 879; Senou et al. 2006b: 463; Koeda et al. 2010a: 75; Koeda et al. 2010b: 81; Hatooka and Yagishita 2013 (in part): 984; Koeda and Motomura 2015: 139; Koeda et al. 2015: 275; Ikeda and Nakabo 2015: 160, figs 4–6; Kaburagi 2016: 99, upper fig. (without scientific name; shown as “Mie-hatampo”); Kimura et al. 2017: 119, fig. 6; Kagoshima City Aquarium Foundation 2018: 210, unnumbered fig.; Koeda 2018b: 298, unnumbered fig.; Koeda 2018c: 342, unnumbered figs; Murase et al. 2019: 132, fig. 285; Koeda 2020a: 409, unnumbered figs; Murase et al. 2021: 166, fig. 341; Koeda 2022: 158, unnumbered fig.; Koeda et al. 2022: 9; Motomura 2023: 128.

Diagnosis

Counts of holotype and non-types are given in Table 1. Dorsal-fin rays VI–VII, 9–10; anal-fin rays III, 40–46; pectoral-fin rays 17–20; pored lateral-line scales 67–78, usually fewer than 73; scale rows above lateral line 8½–10½; scale rows below lateral line 19–22; circumpeduncular scales 24; gill rakers 7–9+19–22 = 28–30; head length 28.1–29.7%; body depth 40.6–44.6%; eye diameter 38.9–43.2%; upper jaw length 50.0–55.6%; maximum 170 mm SL; snout sharp; scales weakly ctenoid, deciduous, thin, semicircular in shape, far wider than long (see Koeda et al. 2013a: fig. 2a); dorsal half of body brown, with golden reflection in fresh specimen; ventral half golden; tip and anterior margin of dorsal fin zonally blackish; faint blackish band on anal-fin base; dusky band on outer edge of anal fin; paired fins pink; posterior half of caudal fin dusky; body pale brown in fixed specimen; black pigmentation on each fins usually persistent, but not in holotype; no blackish blotch on pectoral-fin base; narrow band of villiform teeth in jaws; abdomen cross-sectional outline V-shaped.

Distribution

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. 8).

Remarks

Pempheris sasakii has been commonly confused with P. nyctereutes and has been presumed to be the junior synonym (e.g., Hayashi 1984), even though both species were described by the same first author. Only Tominaga (1963) showed the difference in anal fin coloration (P. sasakii: margin of anal fin fuscous vs P. nyctereutes: margin of anal fin pale) between these two species and described the details of both as valid species. In our morphological observations, however, several P. nyctereutes specimens have dusky margins on the anal fin; thus, this character was not diagnostic for identifying those species. The present comparison based on both species indicated that these two species can be distinguished by the counts of scale rows below lateral line (19–22 in P. sasakii vs 25–27 in P. nyctereutes) with modal difference of pored lateral-line scales (72–81 usually > 79 vs 67–78 usually fewer than 73). The coloration of the species slightly differs in that the former has golden body (sometimes silverish) in fresh condition, but the latter has copper to silver coloration: compare Figs 7, 11). Although the morphological differences between the two species are very few, the molecular analyses strongly supported the intraspecific difference which revealed that sequences of P. nyctereutes and P. sasakii differed by more than 3.1% over mitochondrial 16S ribosomal DNA and COI, comprising different monophyletic groups (Fig. 3).

Figure 11. 

Pempheris sasakii A fresh specimen (KAUM–I. 94368, 126.7 mm SL, Uchinoura Bay, Kimotsuki, Kagoshima) and B underwater photograph (lower: Minamisatsuma, Kagoshima, Japan).

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. 8).

Pempheris schwenkii Bleeker, 1855

Figs 12, 13, Suppl. material 2 Standard Japanese name: Minami-hatampo

Pempheris schwenkii Bleeker, 1855: 314 (type locality: Batu Islands, Sumatera Utara Province, Indonesia); Hatooka 1997 (in part): 380, unnumbered fig. (p. 381, lower middle fig.); Randall and Lim 2000: 622; Hatooka 2002 (in part): 878; Yoshigou and Nakamura 2002: 107; Yoshigou and Nakamura 2003: 49; Yoshigou 2004: 19; Chen 2003: 134, unnumbered fig.; Senou et al. 2006a: 77; Senou et al. 2007: 56; Shao et al. 2008: 254; Yoshino 2008 (in part): 211; Ito 2009: 80, unnumbered fig.; Chen et al. 2010: 265, fig. C, E; Koeda et al. 2010a: 75; Koeda et al. 2010b: 81; Motomura et al. 2010: 131, fig. 252; Shen and Wu 2011: 498, unnumbered fig.; Koeda et al. 2012a: 71; Koeda et al. 2012b: 1086; Miura 2012: 59 (without scientific name; shown as “Minami-hatampo”); Chiang et al. 2014: 183, unnumbered fig.; Hatooka and Yagishita 2013 (in part): 984; Koeda et al. 2013a: 235; Koeda et al. 2013b: 222, fig. 1; Koeda et al. 2013c: 126; Motomura et al. 2013 (in part): 168; Shao et al. 2013 (in part): 163, unnumbered fig. (lower); Koeda et al. 2014: 314; Koeda and Motomura 2015: 139 ; Motomura and Matsuura 2014: 271, unnumbered figs; Koeda et al. 2015: 275; Koeda et al. 2016a: 519; Koeda et al. 2016c: 8, fig. 3H; Koeda and Motomura 2017a; Koeda and Motomura 2017b: 266, fig. 3D, E; Kimura et al. 2017 (in part): 120, fig. 1; Planning and Tourism Division of Kikai Town 2017: 4, unnumbered figs; Nakae et al. 2018: 266; Koeda 2018b: 298, unnumbered fig. (lower right fig.); Koeda 2018b: 194, unnumbered figs; Mochida and Motomura 2018: 30; Koeda 2019: 929, unnumbered figs; Fujiwara and Motomura 2020: 28; Koeda 2020b: 929, unnumbered figs; Motomura and Uehara 2020: 45; Koeda et al. 2022: 10; Motomura 2023: 129 (in part).

Pempheris adusta (not Bleeker, 1877): Shimose 2021: 122, fig. B.

Pempheris japonicus (not Döderlein, 1883): Snyder 1912: 497.

Pempheris oualensis (not Cuvier, 1831): Chen 2003: 134, unnumbered fig.; Shao and Chen 2003: 255, unnumbered figs; Yang et al. 2013: 167, unnumbered fig.

Liopempheris vanicolensis (not Cuvier, 1831): Jordan and Hubbs 1925: 229.

Pempheris vanicolensis (not Cuvier, 1831): Okada 1938: 179; Okada and Matsubara 1938: 179 (in part); Matsubara 1955: 590 (in part); Aoyagi 1948: 49; The Marine Ecological Researching Society of Kagoshima University 1966: 19; The Marine Ecological Researching Society of Kagoshima University 1967: 32; Takahashi 1970: 58; Chen et al. 2010: 266, fig. A; Chang et al 2011: 46.

Pempheris xanthoptera Tominaga, 1963 (in part paratypes): 287; Masuda et al 1975 (in part): 199, pl. 33-C; Yoshino et al. 1975: 75; Hayashi 1984 (in part): 160.

Pempheris sp.: Uchida 1933: 218 (in part).

Diagnosis

Counts of holotype and paratypes are given in Table 2. Dorsal-fin rays VI–VII, very rarely VII, 9–10, very rarely 10; anal-fin rays III, 35–42, usually > 37; pectoral-fin rays 16–18; pored lateral-line scales 44–53; scale rows above lateral line 3½ or 4½, very rarely 4½; scale rows below lateral line 10–13; predorsal scales 23–30; circumpeduncular scales 10–12; gill rakers 6–9+18–22 = 25–30; head length 27.9–33.0%; body depth 40.2–47.2%; eye diameter 35.5–45.5%; upper jaw length 50.0–56.7%; maximum 126 mm SL; scales weakly ctenoid, deciduous, thin, semicircular in shape, far wider than long (see Koeda et al. 2013a: fig. 2a); body golden in day time and silverish in night time; tip of dorsal fin and/or anterior margin of dorsal fin blackish; faint blackish band on anal-fin outer margin; blackish band on anal-fin base; posterior margin of caudal fin dusky; no blackish blotch on pectoral-fin base; posterior nostril usually slit-like; narrow band of villiform teeth in jaws; abdomen cross-sectional outline V-shaped.

Distribution

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. 13). Specimens are collected from the caves or crevasses in coral reef areas of 0–25 m depth.

Figure 12. 

Fresh specimen of Pempheris schwenkii, NMMB-P 27013, 100.3 mm SL, Hengchung, Pingtung, Taiwan.

Figure 13. 

Distribution of Pempheris schwenkii (diamonds) and P. xanthoptera (solid triangles and star for type locality) based on the collection locality of the specimens. Open triangles for literature records of P. xanthoptera.

Remarks

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. Bleeker (1855) described P. schwenkii based on type specimens collected from Batu Island of western Indonesia, eastern Indian Ocean. Although a significant genetic difference is observed among the specimens from these three localities, the morphologies of the species are very similar. Furthermore, the morphological characters were difficult to determine from the dehydrated condition of the syntypes of P. schwenkii (RMNH.PISC.6160). The species composition of genus Pempheris around this area was closer to that of the western Pacific than that of the Indian Ocean. Therefore, P. schwenkii was determined as the name of the Pacific Ocean species (Koeda et al. 2014) and the Indian Ocean species described as a new species, P. tominagai Koeda, Yoshino & Tachihara, 2014. In the present study, a single specimen collected from the Andaman Sea was discovered, and is identified as P. schwenkii based on the pink caudal fin (vs yellow in P. tominagai). This fact supports the conclusion of Koeda et al. (2014) that P. schwenkii may be widely distributed from the Pacific to the Andaman Sea including the type locality (Batu Islands, Sumatera Utara Province, Indonesia) of the species.

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. 13), but the genetic mixability did not appear in the genetic structure analysis (unpublished data); the genetic identification and the diagnostic caudal-fin colorations (pink to brown in P. schwenkii vs yellow in P. xanthoptera) were well matched (Fig. 3).

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 Tominaga (1963) (e.g., Koeda 2017b, 2018a, b, 2020a; Kimura et al. 2017); see remarks of P. xanthoptera]. The Japanese name “Minami-hatampo” was first given by Okada (1938) for P. vanicolensis in his list. His identification may follow Jordan and Hubbs (1925) which indicated that Snyder (1912)’s P. japonica from Okinawa-jima Island was a misidentification of P. vanicolensis. However, P. vanicolensis is very rare in Japanese waters, and has never been collected from Okinawa-jima Island (Nakamura et al. 2022). In addition, the re-examination of the Snyder’s specimen of P. japonica (CAS-SU 22002) revealed that it was in fact a misidentification of P. schwenkii, which is the most common species around Okinawa Island. Although Okada (1938) included Kyushu in the distribution of “Minami-hatampo” which is the range of P. xanthoptera, the situations mentioned above suggest that his species should be P. schwenkii. These facts indicate that the standard Japanese name “Minami-hatampo” should be adopted for the species P. schwenkii.

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 Koeda, Yoshino & Tachihara, 2013

Figs 14, 15, Suppl. material 2 Standard Japanese name: Daito-hatampo

Pempheris ufuagari Koeda et al. 2013a: 232, fig. 1a (type locality: Minamidaito-jima Island, Daito Islands, Japan); Koeda and Motomura 2015: 275; Koeda 2017a: 11, fig. 2 (lower fig.); Koeda and Motomura 2017a; Koeda 2018b: 299, unnumbered fig.

Pempheris oualensis (not Cuvier, 1831): Masuda and Kobayashi 1994 (in part); 180, fig. 5; Hatooka 1997 (in part): 380, unnumbered fig. (p. 381, upper left fig.).

Pempheris otaitensis (not Cuvier, 1831): Randall et al. 1997 (in part): 35, pl. 8, fig. F.

Diagnosis

Counts of holotype and paratypes are given in Table 1 of Koeda et al. (2013a). Dorsal-fin rays VI, 9; anal-fin rays III, 39–43; pectoral-fin rays 17–18; pored lateral-line scales 62–71; scale rows above lateral line 6½–7½ (usually 7½); scale rows below lateral line 15–18; predorsal scales 37–44; circumpeduncular scales 20–22; gill rakers 8–9 + 20–21 = 28–30; head length 26.2–29.9%; body depth 39.7–44.4%; eye diameter 37.5–45.8%; upper jaw length 48.8–53.4%; maximum 197 mm SL, usually < 170 mm SL; scales weakly ctenoid, deciduous, thin, semicircular in shape, far wider than long (Koeda et al. 2013a: see fig. 2a); body silver; distinct blackish blotch on pectoral-fin base; dorsal fin yellow with tip blackish; blackish band on anal-fin outer margin; caudal fin bright yellow with blackish posterior margin; lacking villiform tooth band extending outside lips; abdomen cross-sectional outline V-shaped.

Distribution

Endemic to the Daito and Ogasawara islands (Fig. 15). Specimens collected from the crevasses with strong current in coral reef areas of 0–20 m depth.

Figure 14. 

Pempheris ufuagari A fresh specimen (KAUM–I. 74550, Chichi-jima Island, Ogasawara Islands, 170.3 mm SL, photo taken by K. Kuriiwa) and B underwater photograph (Dobuiso, Ogasawara Islands, Japan).

Figure 15. 

Distribution of Pempheris ufuagari (triangles) and P. vanicolensis (diamonds) based on the collection localities of the specimens.

Remarks

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

Figs 15, 16, Suppl. material 2 Standard Japanese name: Kibire-hatampo

Pempheris vanicolensis Cuvier, 1831: 305 (type locality: Vanikoro Island, Santa Cruz Islands); Shao and Chen 1991: 163, unnumbered fig.; Shao et al. 1992: 177, unnumbered fig.; Shen 1993: 391, pl. 114, fig. 2; Koeda et al. 2010b: 78, fig. 1; Koeda et al. 2012a: 71; Koeda et al. 2013a: 237; Koeda et al. 2013b: 127; Chiang et al. 2014: 183, unnumbered fig.; Koeda et al. 2014: 327; Koeda and Motomura 2015: 139; Koeda et al. 2015: 275; Koeda et al. 2016b: 50, fig. 226; Koeda 2017a: 5; Koeda 2018b: 299, unnumbered fig.; Koeda 2019: 929, unnumbered figs; Koeda et al. 2022: 11; Nakamura et al. 2022: 1, fig. 1.

Pempheris sp.: Shao et al. 2013 (in part): 161, unnumbered fig. (middle fig.); Shao et al. 2008: 254; Hatooka and Yagishita 2013 (in part): 984.

Diagnosis

Counts of holotype and non-types are given in Table 2 of Koeda et al. (2010b). Dorsal-fin rays VI, 9; anal-fin rays III, 38–43; pectoral-fin rays 17–19; pored lateral-line scales 56–65; scale rows above lateral line 5½–6½, usually 5½; scale rows below lateral line 12–15; predorsal scales 30–35; circumpeduncular scales 16–18; gill rakers 8+19–21 = 27–29; head length 29.1–31.7%; body depth 42.4–46.3%; eye diameter 37.5–42.9%; upper jaw length 48.9–55.3%; maximum 156 mm SL; snout rounded; scales weakly ctenoid, deciduous, thin, semicircular in shape, far wider than long (Koeda et al. 2013a: see fig. 2a); body silverish to copperish in fresh specimen collected in day time, but silver in night time; tip of dorsal fin distinctly blackish; usually distinct blackish band on outer edge of anal fin; pectoral fin bright yellow (disappear in fixed specimens) with lacking blackish blotch on its base; posterior margin of caudal fin blackish; narrow band of villiform teeth in jaws; abdomen cross-sectional outline V-shaped.

Figure 16. 

Pempheris vanicolensis A fresh specimen (KAUM–I. 65386, 132.7 mm SL, Hengchung, Pingtung, Taiwan) and underwater photograph (Palau).

Distribution

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. 15). Specimens were collected from 0–2 m depth in Japan but are known from deeper (ca 20 m) in other areas. KK observed a small school of this species at Kuchinoerabu-jima Island (24 Aug. 2016).

Remarks

This species was described by Cuvier (1831), and the name P. vanicolensis has been used for several species, particularly from the Indian Ocean and the Red Sea. However, this species has only been collected from the Pacific Ocean and not from the Indian Ocean as shown in the present study. Jordan and Hubbs (1925) reported that the “P. japonica” documented by Snyder (1912) from Okinawa Island was a misidentification of P. vanicolensis. The early period of modern-day ichthyology in Japan probably followed Jordan and Hubbs (1952), and P. vanicolensis was recognized as “Minami-hatampo” in the species lists of Japanese waters (e.g., Okada 1938; Matsubara 1955). However, P. vanicolensis is very rare in Japan, and re-examination of Snyder’s specimen (SU 22002) revealed that it was a misidentification of P. schwenkii, which is the most common species around Okinawa Island.

Pempheris xanthoptera Tominaga, 1963

Figs 13, 17, Suppl. material 2 New standard Japanese name: Mizuho-hatampo

Pempheris xanthoptera Tominaga, 1963: 286, fig. 12 (type locality: Manazuri, Kanagawa Prefecture, Japan); Hiyama and Yasuda 1971: 204, fig. 440; Masuda et al. 1975: 199, pl. 33-C; Hayashi 1984: 160, pl. 151-D; Kohno 1986: 135, fig. 1; Randall et al. 1997: 35; Koeda et al. 2022: 11.

Pempheris molucca (not Cuvier, 1829): Temminck and Schlegel 1844: 85, pl. 44, fig. 3.

Pempheris oualensis (not Cuvier, 1831): Mochizuki 1995: 389, unnumbered fig. (in part).

Pempheris schwenkii (not Bleeker, 1855): Masuda and Kobayashi 1994: 180, fig. 6; Mochizuki 1995: 389, unnumbered fig.; Hatooka 1997 (in part): 380, unnumbered fig. (p. 381, lower left fig.); Hatooka 2002 (in part): 878; Takayama et al. 2003: 1317, fig. 2; Senou et al. 2006b: 463; Aramata 2007: 172, unnumbered figs; Yoshino 2008 (in part): 211; Takagi et al. 2010: 69, unnumbered figs; Kohno et al. 2011: 208, unnumbered fig.; Senou et al. 2012: 212; Hatooka and Yagishita 2013 (in part): 984; Koeda et al. 2013a: 237; Motomura et al. 2013 (in part): 168, unnumbered fig.; Kawano et al. 2014: 48; Koeda et al. 2014: 327; Ikeda and Nakabo 2015: 160, figs 1–3; Takeuchi et al. 2015: 8; Iwatsubo et al. 2016: 22, unnumbered figs; Kaburagi 2016: 98, lower fig. (without scientific name; indicated as “Minami-hatampo”); Kimura et al. 2017 (in part): 120, fig. 2; Koeda 2017a: 9, fig. 2 (middle); Koeda 2017b: 190, unnumbered fig.; Koeda and Motomura 2017a; Kagoshima City Aquarium Foundation 2018: 210, unnumbered fig.; Koeda 2018b: 298, unnumbered fig. (lower left fig.); Koeda 2018c: 343, unnumbered figs; Murase et al. 2019: 132, fig. 284; Koeda 2020a: 407, unnumbered figs; Murase et al. 2021: 166, fig. 340; Koeda 2022: 158, unnumbered fig.; Motomura 2023: 129 (in part).

Pempheris vanicolensis (not Cuvier, 1831): Okada 1938: 179 (in part); Okada and Matsubara 1938: 179 (in part); Matsubara 1955: 590 (in part); Abekawa and Nishi 1969: 24.

Pempheris japonica (not Döderlein, 1883): Nakamura 1993: 148, fig. 6.

Diagnosis

Counts of of holotype and paratypes are given in Table 2. Dorsal-fin rays VI–VII, 9; anal-fin rays III, 35–42; pectoral-fin rays 16–19; pored lateral-line scales 45–54; scale rows above lateral line 3 1/2; scale rows below lateral line 10–13; predorsal scales 23–28; circumpeduncular scales 12–14; gill rakers 7–9+18–21 = 25–27; head length 27.9–31.9%; body depth 39.3–45.4%; eye diameter 36.4–44.1%; upper jaw length 48.3–57.1%; maximum 137 mm SL; scales weakly ctenoid, very deciduous, thin, semicircular in shape, far wider than long (see Koeda et al. 2013a: fig. 2a); body golden in daytime, but silver in night time; no blackish blotch on; tip of dorsal fin distinctly blackish; anal-fin base zonal blackish, and margin very faintly blackish; caudal fin yellow (disappears in fixed specimens); pectoral and pelvic fin hyaline or pink; posterior margin of caudal fin dusky; body light brown to dark brown in fixed specimen; black pigmentation on each fins usually persistent; no blackish blotch on pectoral-fin base; posterior nostril usually open, not compressed; narrow band of villiform teeth in jaws; abdomen cross-sectional outline V-shaped.

Figure 17. 

Fresh specimen of Pempheris xanthoptera, KBF-I 00268, 123.0 mm SL, Amaji, Otsuki, Kochi.

Distribution

Endemic to the Northwest Pacific, recorded only from Japan, Jeju Island in Korea (Kim and Sakai 2004), and China. In Japanese waters, this species is known from Pacific coast (north to Boso Peninsula in Chiba Prefecture, south to Kagoshima Prefecture), Tsushima Island, East China Sea coast (north to Goto Islands in Nagasaki Prefecture, south to Kagoshima Prefecture), Izu-oshima, Miyake-jima and Hachijo-jima islands in Izu Islands, Chichi-jima, Haha-jima, Ototo-jima islands in Ogasawara Islands, Tanega-shima, Yaku-shima, Kuchinoerabu-jima, Iou-jima and Take-shima islands in northern Ryukyu Archipelago (Fig. 13).

Remarks

Although Tominaga (1963) described P. xanthoptera based on its differences in fin color and distributional pattern from P. schwenkii, the former has been usually considered as a junior synonym of the latter in recent publications in Japan (see the synonym list) without any discussion. Our genetic analysis revealed apparent differences between these two species with high node support values (Fig. 3). The morphological comparison showed the additional small difference between these two species, such as P. xanthoptera has the posterior nostril usually open, not compressed (vs slit-like; see Koeda et al. 2014: fig. 8), and caudal fin yellow (vs pink to brown). The validity of P. xanthoptera was discussed in the remarks of P. schwenkii.

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.

Acknowledgments

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.

Additional information

Conflict of interest

The authors have declared that no competing interests exist.

Ethical statement

No ethical statement was reported.

Funding

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).

Author contributions

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.

Author ORCIDs

Keita Koeda https://orcid.org/0000-0003-3932-3002

Manabu Bessho-Uehara https://orcid.org/0000-0002-7388-464X

Data availability

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

References

  • Abekawa Y, Nishi G (1969) Fish fauna of Hanaze, Ibusuki-gun, Kagoshima Prefecture. KYANOΣ OIKΣ 13: 23–27. [In Japanese]
  • Aoyagi H (1948) Study on coral reef fishes from the Ryukyu Archipelago II, Synodontidae, Plotosidae, Pempheridae, and others. Zoological Magazine 58: 48–50. [In Japanese]
  • Aramata H (2007) The Aramata atlas of wonder fishes. Shinshokan, Tokyo, 390 pp. [In Japanese]
  • Bleeker P (1855) Bijdrage tot de kennis der ichthyologische fauna van de Batoe Eilanden. Natuurkundig Tijdschrift voor Nederlandsch-Indië 8: 305–328.
  • Bleeker P (1877) Révision des espèces de Pempheris de l’inde archipélagique. Archives Néerlandaises des Sciences Exactes et Naturelles 12: 42–54.
  • Chang C-W, Wang S-I, Yang C-J, Shao K-T (2011) Fish fauna in subtidal waters adjacent to the National Museum of Marine Biology and Aquarium. Platax 8: 41–45.
  • Chen C-H (2003) Fishes of Penghu. Fisheries Research Institute, Council of Agriculture, Keelung, [xxxvi +] 379 pp. [In Chinese]
  • Chen J-P, Shao K-T, Lin C-P (1995) A checklist of reef fishes from Tungsha Tao (Pratas Island), South China Sea. Acta Zoologica Taiwanica 6: 13–40.
  • Chen J-P, Shao K-T, Jan R-Q, Kuo J-W, Chen J-Y (2010) Marine fishes in Kenting National Park (1st revised ed.). Kenting National Park Headquarters, Hengchun, 656 pp. [In Chinese]
  • Chiang W-C, Lin P-L, Chen W-Y, Liu D-C (2014) Marine Fishes in Eastern Taiwan. Fisheries Research Institute, Keelung, [x +] 337 pp. [In Chinese]
  • Cuvier G (1829) Le Règne Animal, distribué d’après son organisation, pour servir de base à l’histoire naturelle des animaux et d’introduction à l’anatomie comparée. 2nd edn. Tome 2. Règne Animal. Poissons. Deterville, Paris, [xv +] 406 pp. https://doi.org/10.5962/bhl.title.1964
  • Cuvier G (1831) Des Pemphérides. In: Cuvier G, Valenciennes A (Eds) Historie Naturelle des Poissons, Vol. 7. F. G. Levraut, Paris-Strasbourg, [xx–xxi+]296–309 [191 pls].
  • Döderlein L (1883) Kaiserlichen Akademie der Wissenschaften in Wien, Sitzung der mathematisch-naturwissenschaftlichen classe vom 14. Juni 1883. In: Steindachner F, Döderlein L (Eds) Beiträge zur Kenntniss der Fische Japan’s (II. ). Anzeiger der Kaiserlichen Akademie der Wissenschaften, Wien, Mathematisch-Naturwissenschaftliche Classe 20 (15), 123–130.
  • Fowler HW (1906) New, rare or little known scombroids, No. 3. Proceedings of the Academy of Natural Sciences of Philadelphia 58: 114–122.
  • Fujiwara K, Motomura H (2020) An annotated checklist of marine and freshwater fishes of Kikai Island in the Amami Islands, Kagoshima, southern Japan, with 259 new records. Bulletin of the Kagoshima University Museum 14: 1–73.
  • Fujiwara K, Tanoue H, Mohri M, Kamano T, Hata K, Okada S, Nagai S, Motomura H (2018) Fishes of the Hibiki-nada Sea and Mi-shima island, Yamaguchi Prefecture, Japan. Journal of National Fisheries University 66: 47–80. [In Japanese with English Abstract]
  • Golani D, Ben-Tuvia A (1986) New records of fishes from the Mediterranean coast of Israel including Red Sea immigrants. Cybium 10: 285–291.
  • Golani D, Diamant A (1991) Biology of the sweeper, Pempheris vanicolensis Cuvier & Valenciennes, a Lessepsian migrant in the eastern Mediterranean, with a comparison with the original Red Sea population. Journal of Fish Biology 38: 819–827. https://doi.org/10.1111/j.1095-8649.1991.tb03621.x
  • Guindon S, Dufayard JF, Lefort V, Anisimova M, Hordijk W, Gascuel O (2010) New algorithms and methods to estimate maximum-likelihood phylogenies: assessing the performance of PhyML 3.0. Systems Biology 59: 307–321. https://doi.org/10.1093/sysbio/syq010
  • Hatooka K (1997) Pempheridae. In: Okamura O, Amaoka K (Eds) Sea fishes of Japan. Yama-kei Publishers co., Ltd., Tokyo, 380–381. [In Japanese]
  • Hatooka K (2000) Pempheridae. In: Nakabo T (Ed.) Fishes of Japan with pictorial keys to the species, second edition. Tokai University Press, Tokyo, 878–879, 1570–1571.
  • Hatooka K (2002) Pempheridae. In: Nakabo T (Ed.) Fishes of Japan with pictorial keys to the species, English edition. Tokai University Press, Tokyo, 878–879, 1562.
  • Hatooka K, Yagishita N (2013) Pempheridae. In: Nakabo T (Ed.) Fishes of Japan with pictorial keys to the species, 3rd edn. Tokai University Press, Tokyo, 983–984, 2020–2021. [In Japanese]
  • Hayashi Y (1984) Pempheridae. In: Masuda H, Amaoka K, Araga C, Uyeno T, Yoshino T (Eds) The fishes of the Japanese Archipelago. Tokai University Press, Tokyo, 160–161 [pls 151, 350]. [In Japanese]
  • Hiyama Y, Yasuda T (1971) Living fishes of the Japanese coastal waters. Kodansha, Tokyo, 337 pp. [In Japanese]
  • Honda H (1972) Fishes of Zamami-jima island. KYANOΣ OIKΣ 15: 41–48. [In Japanese]
  • Ikeda H, Nakabo T (2015) Fishes of the Pacific coasts of southern Japan. Tokai University Press, Hadano, 597 pp. [In Japanese]
  • Ito K (2009) The underwater in Okinawa. Data house, Tokyo, 457 pp. [In Japanese]
  • Jordan DS, Hubbs LC (1925) Record of the fishes obtained by David Starr Jordan in Japan, 1922. Mem Carnegie Mus 10(2): 93–346, [pls 5–12]. https://doi.org/10.5962/p.234844
  • Jordan DS, Tanaka S, Snyder JO (1913) A catalogue of the fishes of Japan. The Journal of the College of Science, Imperial University of Tokyo 33: 1–497.
  • Kaburagi K (2016) Field guide to fishes of Tanega-shima Island collected by line and hook. Tamashidasha, Nishinoomote, 158 pp. [In Japanese]
  • Kagoshima City Aquarium Foundation (2018) Fishes collected with set nets in Kagoshima and confirmed by Kagoshima City Aquarium, 2nd edn. Kagoshima City Aquarium Foundation, Kagoshima, 335 pp. [In Japanese]
  • Katoh K, Standley DM (2013) MAFFT multiple sequence alignment software version 7: improvements in performance and usability. Molecular Biology and Evolution 30: 772–780. https://doi.org/10.1093/molbev/mst010
  • Kawano M, Doi H, Hori S (2014) List of the fishes in the southwestern Japan Sea off Yamaguchi Prefecture. Bulletin of Yamaguchi Prefecture Fisheries Research Center 11: 1–30. [In Japanese with English abstract]
  • Kim B-J, Sakai K (2004) On the occurrence of Pempheris schwenkii (Pempheridae) at Jeju Island: an addition to Korean ichthyofauna. Korean Journal of Ichthyology 16(2): 181–184.
  • Kimura Y, Hibino Y, Miki R, Minetoma T, Koeda K (2017) Field guide to fishes of Kuchinoerabu-jima Island in the Osumi Group, Kagoshima, southern Japan. The Kagoshima University Museum, Kagoshima, 200 pp. [In Japanese]
  • Klunzinger CB (1879) Die von Müller’sche Sammlung Australischer Fische. Anzeiger der Österreichische Akademie der Wissenschaften 16: 254–261.
  • Koeda K (2017a) Fishes of family Pempheridae (Teleostei: Perciformes) of the Ogasawara Islands, Japan. Bulletin of the Biogeographical Society of Japan 71: 1–13. [In Japanese with English abstract]
  • Koeda K (2017b) Pempheridae. In: Iwatsubo H, Motomura H (Eds) Field guide to fishes of Kagoshima Bay in southern Kyushu, Japan. Kagoshima Museum of Aquatic Biodiversity, Kagoshima and the Kagoshima University Museum, Kagoshima, 189–190. [In Japanese]
  • Koeda K (2018a) Pempheridae. In: Motomura H, Hagiwara K, Senou H, Nakae M (Eds) Identification guide to fishes of the Amami Islands, Japan. Kagoshima University Museum, Kagoshima, Yokosuka City Museum, Yokosuka, Kanagawa Prefectural Museum of Natural History, Odawara, and National Museum of Nature and Science, Tsukuba, 193–194. [In Japanese]
  • Koeda K (2018b) Pempheridae. In: Nakabo T (Ed.) The natural history of the fishes of Japan. Shogakukan, Tokyo, 298–299. [In Japanese]
  • Koeda K (2018c) Pempheridae. In: Koeda K, Hata H, Yamada M, Motomura H (Eds) Field guide to fishes landed at Uchinoura Fishing Port, Kagoshima, Japan. The Kagoshima University Museum, Kagoshima, 339–343. [In Japanese]
  • Koeda K (2018d) Pempheridae. In: Kimura S, Imamura H, Nguyen VQ, Pham TD (Eds) Fishes of Ha Long Bay the World the world natural heritage site in northern Vietnam. Fisheries Research Laboratory, Mie University, Shima, Japan, 195–196.
  • Koeda K (2019) Family Pempheridae. In: Koeda K, Ho H-C (Eds) Fishes of southern Taiwan. National Museum of Marine Biology & Aquarium, Pingtung, Taiwan, 925–930.
  • Koeda K (2020a) Pempheridae. In: Koeda K, Hata H, Yamada M, Motomura H (Eds) Fishes from markets in Osumi Peninsula, Kagoshima, Japan. The Kagoshima University Museum, Kagoshima, 406–411. [In Japanese]
  • Koeda K (2020b) Family Pempheridae. In: Koeda K, Ho H-C (Eds) Fishes of southern Taiwan, 2nd edn. National Museum of Marine Biology & Aquarium, Pingtung, Taiwan, 925–930.
  • Koeda K (2022) Family Pempheridae. In: Iwatsubo H, Ito M, Yamada M, Motomura H (Eds) Field guide to fishes of the East China Sea side of Satsuma Peninsula in Kagoshima, southern Kyushu, Japan. Kagoshima Museum of Aquatic Biodiversity, Kagoshima and the Kagoshima University Museum, Kagoshima, 158–158.
  • Koeda K, Motomura H (2015) First records of Pempheris adusta (Perciformes: Pempheridae) from Kuchierabujima, Nakano-shima, and Tokuno-shima islands in the Satsunan Islands and the Kagoshima mainland, southern Japan with some biological comments. Nature of Kagoshima 41: 139–144. [In Japanese]
  • Koeda K, Motomura H (2017a) A new species of Pempheris (Perciformes: Pempheridae) endemic to the Ogasawara Islands, Japan. Ichthyological Research. https://doi.org/10.1007/s10228-017-0586-3 [also appeared in Ichthyological Research 65: 21–28]
  • Koeda K, Motomura H (2017b) Annotated list of fish specimens taken from fish stomach deposited at the Kagoshima University Museum, Japan. Nature of Kagoshima 43: 257–269. [In Japanese]
  • Koeda K, Imai H, Yoshino T, Tachihara K (2010a) First and northernmost record of Pempheris oualensis (Pempheridae), from Ryukyu Archipelago, Japan. Biogeography 12: 71–75.
  • Koeda K, Yoshino T, Imai H, Tachihara K (2010b) Description of new Japanese and northernmost records of pempherid fish, Pempheris vanicolensis, from Iriomote Island, southern Ryukyu Archipelago. Biogeography 12: 77–82.
  • Koeda K, Yoshino T, Tachihara K (2012a) First and southernmost records of Pempheris japonica (Pempherididae) from Okinawa Island with the description of juvenile growth. Bulletin of the Biogeographical Society of Japan 67: 65–73. [In Japanese with English abstract]
  • Koeda K, Ishihara T, Tachihara K (2012b) The reproductive biology of Pempheris schwenkii (Pempheridae) on Okinawa Island, southwestern Japan. Zoological Studies 51: 1086–1093.
  • Koeda K, Yoshino T, Tachihara K (2013a) Pempheris ufuagari sp. nov., a new species in the genus Pempheris (Perciformes, Pempheridae) from the oceanic islands of Japan. Zootaxa 3609: 231–238. https://doi.org/10.11646/zootaxa.3609.2.9
  • Koeda K, Yoshino T, Tachihara K (2013b) Identificational keys of Pempheris adusta Bleeker, 1877 (Pempheridae) with comments on its standard Japanese name. Japanese Journal of Ichthyology 60: 123–128. [In Japanese with English abstract]
  • Koeda K, Fukagawa T, Ishihara T, Tachihara K (2013c) Reproductive biology of nocturnal reef fish Pempheris adusta (Pempheridae) in Okinawa Island, Japan. Galaxea 15: 221–228. https://doi.org/10.3755/galaxea.15.221
  • Koeda K, Yoshino T, Imai H, Tachihara K (2014) A review of the genus Pempheris (Perciformes, Pempheridae) of the Red Sea, with description of a new species. Zootaxa 3793: 301–330. https://doi.org/10.11646/zootaxa.3793.3.1
  • Koeda K, Kaburagi K, Motomura H (2015) Records of Pempheris oualensis (Perciformes: Pempheridae) from the Satsunan Islands, Ryukyu Archipelago, Japan. Bulletin of the Biogeographical Society of Japan 70: 275–282. [In Japanese with English abstract]
  • Koeda K, Ishihara T, Fukagawa T, Tachihara K (2016a) Life cycle differences between two species of genus Pempheris based on age determination around Okinawa-jima Island. Ichthyological Research 63: 519–528 https://doi.org/10.1007/s10228-016-0526-7
  • Koeda K, Hibino Y, Yoshida T, Kimura Y, Miki R, Kunishima T, Sasaki D, Furukawa T, Sakurai M, Eguchi K, Suzuki H, Inaba T, Uejo T, Tanaka S, Fujisawa M, Wada H, Uchiyama T (2016b) Annotated checklist of fishes of Yonagunijima Island, the westernmost island in Japan. The Kagoshima University Museum, Kagoshima, 119 pp.
  • Koeda K, Fujii T, Koeda S, Motomura H (2016c) Fishes of Yoro-jima and Uke-jima islands in the Amami Islands: 89 new specimen-based records. Memories of Facility of Fisheries Kagoshima University 65: 1–20.
  • Koeda K, Aizawa M, Sakamoto K, Ueshima R (2022) Report on specimens of the family Pempheridae (Teleostei: Perciformes) deposited in the Department of Zoology, The University Museum, The University of Tokyo. The University Museum, The University of Tokyo Material Reports 128: 1–16.
  • Kohno H (1986) Larvae and juveniles of pempherid fishes, Pempheris xanthoptera and P. japonica. Japanese Journal of Ichthyology 33: 135–144. https://doi.org/10.1007/BF02905841
  • Kohno H, Konou K, Tokoo T (2011) A photographic guide to the fishes in Tokyo Bay. Heibonsha, Tokyo, 374 pp. [In Japanese]
  • Kurata T, Mimura T, Kusadokoro K (1971) Fish fauna of the Ogasawara Islands and the catch tendencies of commercially important fishes. Report of the Tokyo Metropolitan Fisheries Experiment Station 216: 1–38. [In Japanese]
  • Lee K-L (1996) Field guide to common fishes of Taiwan 2. Deep sea demersal fishes. Outdoor Life Books Co. , Ltd., Taipei, 188 pp. [In Chinese]
  • Masuda H, Kobayashi Y (1994) Grand atlas of fish life modes. Tokai University Press, Tokyo, 379 pp. [In Japanese]
  • Masuda H, Araga C, Yoshino T (1975) Coastal fishes of southern Japan. Tokai University Press, Tokyo, 379 pp. [In Japanese]
  • Matsubara K (1955) Fish morphology and hierarchy. Ishizaki-shoten, Tokyo, 789 pp. [In Japanese]
  • Miura N (2012) Churaumi Ichiba Zukan, Chinen Ichiba no Sakana tachi [Fishes at Chinen Market, Okinawa]. Wave Kikaku, Yonabaru, 140 pp. [In Japanese]
  • Mochida I, Motomura H (2018) An annotated checklist of marine and freshwater fishes of Tokunoshima island in the Amami Islands, Kagoshima, southern Japan, with 214 new records. Bulletin of the Kagoshima University Museum 10: 1–80.
  • Mochizuki K (1995) Pempheridae. In: Konishi H (Ed.) Fishes-new color guide for sportfishermen. Weekly Sunday Fishing Co. Ltd, Osaka, 389–389.
  • Mooi RD (1998) A new species of the genus Pempheris (Teleostei: Pempherididae) from Rapa Iti, French Polynesia. Pacific Science 52: 154–160.
  • Mooi RD, Jubb RN (1996) Description of two new species of the genus Pempheris (Pisces: Pempherididae) from Australia, with a provisional key to Australian species. Records of the Australian Museum 45: 117–130. https://doi.org/10.3853/j.0067-1975.48.1996.284
  • Motomura H (2023) An annotated checklist of marine and freshwater fishes of Tanega-shima amd Mage-shima islands in the Osumi Islands, Kagoshima, southern Japan, with 536 new records. Bulletin of the Kagoshima University Museum 20: 1–250.
  • Motomura H, Matsuura K (2014) Field guide to fishes of Yoron Island in the middle of the Ryukyu Islands, Japan. The Kagoshima University Museum, Kagoshima and the National Museum of Nature and Science, Tsukuba, 648 pp.
  • Motomura H, Uehara K (2020) An annotated checklist of marine and freshwater fishes of Okinoerabu Island in the Amami Islands, Kagoshima, southern Japan, with 361 new records. Bulletin of the Kagoshima University Museum 12: 1–125.
  • Motomura H, Kuriiwa K, Katayama E, Senou H, Ogihara G, Meguro M, Matsunuma M, Takata Y, Yoshida T, Yamashita M, Kimura S, Endo H, Murase A, Iwatsuki Y, Sakurai Y, Harazaki S, Hidaka K, Izumi H, Matsuura K (2010) Annotated checklist of marine and estuarine fishes of Yaku-shima Island, Kagoshima, southern Japan. In: Motomura H, Matsuura K (eds) Fishes of Yaku-shima Island – A World Heritage island in the Osumi Group, Kagoshima Prefecture, southern Japan. National Museum of Nature and Science, Tokyo, 65–247.
  • Motomura H, Dewa S, Furuta K, Matsuura K (2013) Fishes of Iou-jima and Take-shima islands, Mishima, Kagoshima, Japan. The Kagoshima University Museum, Kagoshima and the National Museum of Nature and Science, Tsukuba, 390 pp.
  • Mouneimne N (1979) Poissons nouveaux pour les côtes Libanaises. Cybium 6: 105–110. [in French]
  • Müller J, Troschel FH (1848) In: Schomburgk RH (Ed.) The history of Barbados; comprising a geographical description of the island and an account of its geology and natural productions. London. The history of Barbados; comprising a geographical description of the island and an account of its geology and natural productions, [xx +] 722 pp [17 pls].
  • Murase A, Miki R, Wada M, Senou H (2019) Coastal and market fishes around Kadogawa Bay, northern part of Miyazaki Prefecture, southern Japan. Nobeoka Marine Science Station, Field Science Center, University of Miyazaki, Nobeoka, 207 pp. [In Japanese]
  • Murase A, Ogata Y, Yamasaki Y, Miki R, Wada M, Senou H (2021) Coastal, shelf and deep-sea fishes around Kadogawa Bay, northern part of Miyazaki Prefecture, southern Japan. Nobeoka Marine Science Station, Field Science Center, University of Miyazaki, Nobeoka, 357 pp. [In Japanese]
  • Nakae M, Motomura H, Hagiwara K, Senou H, Koeda K, Yoshida T, Tashiro S, Jeong B, Hata H, Fukui Y, Fujiwara K, Yamakawa T, Aizawa M, Shinohara G, Matsuura K (2018) An Annotated Checklist of Fishes of Amami-oshima Island, the Ryukyu Islands, Japan. Memoirs of the National Science Museum (Tokyo) 52: 205–361.
  • Nakamura H (1933) Juveniles of reef fishes appeared around Kominato (1). Yoshokukaishi 3: 145–148.
  • Nakamura J, Koeda K, Motomura H (2022) Northernmost record of Pempheris vanicolensis (Perciformes: Pempheridae) from Kikai-jima Island, Amami Islands, Japan. Japanese Journal of Ichthyology. https://doi.org/10.11369/jji.22-004 [also appeared in Japanese Journal of Ichthyology 69: 219–223]
  • Ogilby JD (1913) Edible fishes of Queensland. Part I. Family Pempheridae. Part II. The gadopseiform percoids. Memoirs of the Queensland Museum 2: 60–80 [pls 18–20].
  • Okada Y (1938) A catalogue of vertebrates of Japan. Maruzen, Co. Ltd, Tokyo, [iv +] 412 pp.
  • Okada Y, Matsubara K (1938) Key to the fishes and fish-like animals of Japan. Sanseido, Tokyo.
  • Planning and Tourism Division of Kikai Town (2017) Traditional island’s fish “Ufumi”. Kikai Publication Relations 604: 4–6. [In Japanese]
  • Randall JE, Lim KKP (2000) A checklist of the fishes of the South China Sea. Raffles Bulletin of Zoology 8: 569–667.
  • Randall JE, Victor BC (2014) Four new fishes of the genus Pempheris (Perciformes: Pempheridae) from the western Indian Ocean. Journal of the Ocean Science Foundation 12: 61–83.
  • Randall JE, Victor BC (2015) Descriptions of thirty-four new species of the fish genus Pempheris (Perciformes: Pempheridae), with a key to the species of the western Indian Ocean. Journal of the Ocean Science Foundation 18: 1–77.
  • Randall JE, Ida H, Kato K, Pyle RL, Earle JL (1997) Annotated checklist of the inshore fishes of the Ogasawara Islands. National Museum of Nature and Science Monographs 11: 1–74.
  • Randall JE, Victor BC, Alpermann TJ, Bogorodsky SV, Mal AO, Satapoomin U, Bineesh KK (2014) Rebuttal to Koeda et al. (2014) on the Red Sea fishes of the perciform genus Pempheris. Zootaxa 3887: 377–392. https://doi.org/10.11646/zootaxa.3887.3.5
  • Sakurai S, Masuda Y, Nagaoka I, Tokioka S, Togashi H (2024) Twenty-nine southern fish species, including species representing the northernmost records, collected around the Oshika Peninsula, Miyagi Prefecture, northern Japan, during October 2023 to February 2024 under abnormally high water temperature. Ichthy, Natural History of Fishes of Japan 45: 68–84. [In Japanese with English abstract]
  • Schmidt PJ (1913) Fishes of the Riu-kiu Islands. Transactions of the Pacific Committee of the Academy of Science of the USSR 1: 19–156 [pls 1–6].
  • Senou H, Shinohara G, Matsuura K, Furuse K, Kato S, Kikuchi T (2002) Fishes of Hachijo-jima Island, Izu Islands Group, Tokyo, Japan. Memoirs of the National Science Museum (Tokyo) 38: 195–237.
  • Senou H, Kodato H, Nomura T, Yunokawa K (2006a) Coastal fishes of the Ie-jima Island, the Ryukyu Islands, Okinawa, Japan. Bulletin of the Kanagawa Prefectural Museum (Natural Science) 35: 67–92.
  • Senou H, Matsuura K, Shinohara G (2006b) Checklist of fishes in the Sagami Sea with zoogeographical comments on shallow water fishes occurring along the coastlines under the influence of Kuroshio Current. Memoirs of the National Science Museum (Tokyo) 41: 389–542.
  • Senou H, Kobayashi Y, Kobayashi N (2007) Coastal fishes of the Miyako Group, the Ryukyu Islands, Japan. Bulletin of the Kanagawa Prefectural Museum (Natural Science) 36: 47–74.
  • Shao K-T, Chen L-W (1991) Field guide to observe nature of Taiwan. Reef angling fish (1). Holiday Press, Taipei, 235 pp.
  • Shao K-T, Chen J-P, Shen S-C (1992) Marine fishes of the Ken-ting National Park. Ken-Ting National Park Headquarters Construction and Planning Administration, Kenting, 427 pp.
  • Shao K-T, Ho H-C, Lin P-L, Lee P-F, Lee M-Y, Tsai C-Y, Liao Y-C, Lin Y-C (2008) A checklist of the fishes of southern Taiwan, northern South China Sea. The Raffles Bulletin of Zoology supplement 19: 233–271.
  • Shao K-T, Shao I-T, Lin P-L (2013) Coral reef fishes of Taiwan. Intimate. , LTD., Taichung, 335 pp.
  • Shaw G (1790) Doubtful, or compressed Sparus. In: White J (Ed.) Journal of a voyage to New South Wales with sixty-five plates of non descript animals, birds, lizards, serpents, curious cones of trees and other natural productions. J.Debrett. Piccadilly, London, 267 pp. [pl. 53]
  • Shen S-C (1984) Coastal fishes of Taiwan. National Taiwan Museum, Taipei, 191 pp. [in Chinese and English]
  • Shen S-C (1993) Fishes of Taiwan. National Taiwan University, Taipei, 960 pp.
  • Shen S-C, Wu K-Y (2011) Fishes of Taiwan. National Museum of Marine Biology & Aquarium, Checheng, 896 pp.
  • Shimose T (2021) Commercial Fishes and shellfishes of Okinawa. Okinawa times, Naha, 207 pp. [In Japanese]
  • Snyder JO (1911) Japanese shore fishes collected by the United States Bureau of Fisheries steamer ‘‘Albatross’’ expedition of 1906. Proceedings of the United States National Museum 42: 399–450 [pls 51–61]. https://doi.org/10.5479/si.00963801.42-1909.399
  • Sonoyama T, Ogimoto K, Hori S, Uchida Y, Kawano M (2020) An annotated checklist of marine fishes of the Sea of Japan off Yamaguchi Prefecture, Japan, with 74 new records. Bulletin of the Kagoshima University Museum 11: 1–152.
  • Takagi M, Hirata T, Hirata S, Nakata C (2010) Fishes of Ainan Ehime. Soufusha shuppan, Matsuyama, 250 pp.
  • Takahashi S (1970) Fish fauna of coral reef of Yoro-jima island. KYANOΣ OIKΣ 14: 57–62.
  • Takayama M, Onuki A, Yoshino T, Yoshimoto M, Ito H, Kohbara J, Somiya H (2003) Sound characteristics and the sound producing system in silver sweeper, Pempheris schwenkii (Perciformes: Pempheridae). Journal of the Marine Biological Association of the UK 83: 1317–1320. https://doi.org/10.1017/S0025315403008750
  • Takemura Y, Yasuta T (1965) The sea fishes of Japanese coastal waters. Hokuryukan, Tokyo, 211 pp.
  • Takeuchi N, Senou H, Seino H (2015) Fish fauna of Tsushima Island, Nagasaki Prefecture, Japan -researches from 1948 to 2015. Bulletin of the Biogeographical Society of Japan 70: 1–11.
  • Tanaka S (1931) On the distribution of fishes in Japanese waters. Journal of the Faculty of Science, Imperial University of Tokyo, Section IV, Zoology 3: 3–90 [3 pls].
  • The Marine Ecological Researching Society of Kagoshima University (1966) Special report of Republic of China. KYANOΣ OIKΣ 9: 1–60.
  • The Marine Ecological Researching Society of Kagoshima University (1967) Report of Yakushima Island. KYANOΣ OIKΣ X: 1–37. [In Japanese]
  • Tominaga Y (1963) A revision of the fishes of the family Pempheridae of Japan. Journal of the Faculty of Science, the University of Tokyo 10: 269–290.
  • Tominaga Y (1968) Internal morphology, mutual relationships and systematic position of the fishes belonging to the family Pempheridae. Japanese Journal of Ichthyology 15: 43–95.
  • Toyama N (1937) Flora and fauna of waters around Ogasawara Islands. Tokyo Metropolitan Government, Tokyo, 93 pp.
  • Trifinopoulos J, Nguyen LT, von Haeseler A, Minh BQ (2016) W-IQ-TREE: a fast online phylogenetic tool for maximum likelihood analysis. Nucleic Acids Research 44: W232–W235. https://doi.org/10.1093/nar/gkw256
  • Uchida K (1933) Life history of Pempheris japonicus, and early development of congeners. Zoological Science 45: 207–218. [In Japanese]
  • Yang C-J, Yang C-M, Li J-J (2013) A Comprehensive Guide to Diving on Orchid Island. National Museum of Marine Biology and Aquarium, Checheng, 359 pp.
  • Yoshigou H (2004) The fishes found from tide-pools and surge sublittoral zone in the Minami-daito (South-Borodino) Island, Japan. Hiwa Museum for Natural History Material Reports 43: 1–51 [10 pls]. [In Japanese with English abstract]
  • Yoshigou H, Nakamura S (2002) Catalogue of fish specimens preserved in Hiwa Museum for Natural History II. Hiwa Museum for Natural History Material Reports 3: 85–136 [pl. 1]. [In Japanese with English abstract]
  • Yoshigou H, Nakamura S (2003) Catalogue of fish specimens preserved in Hiwa Museum for Natural History III. Hiwa Museum for Natural History Material Reports 4: 29–75 [pl. 1]. [In Japanese with English abstract]
  • Yoshigou H, Naito J, Nakamura S (2001) Catalogue of fish specimens preserved in Hiwa Museum for Natural History. Hiwa Museum for Natural History Material Reports 2: 119–168. [In Japanese with English abstract]
  • Yoshino Y (2008) Marine fishes of Japan. Yama-kei Publishers co. , LTD, Tokyo, 543 pp.
  • Yoshino T, Nishijima S, Shinohara S (1975) Catalogue of fishes of the Ryukyu Islands. Bulletin of Science and Engineering Division University of the Ryukyus; Mathematics & Natural Sciences 20: 61–118.
  • Zama A, Fujita K (1977) An annotated list of the fishes from the Ogasawara Islands. Journal of the Tokyo University of Fisheries 63: 87–138.

Supplementary materials

Supplementary material 1 

Preserved specimens of genus Pempheris examined in the present study

Keita Koeda, Manabu Bessho-Uehara

Data type: xlsx

This dataset is made available under the Open Database License (http://opendatacommons.org/licenses/odbl/1.0/). The Open Database License (ODbL) is a license agreement intended to allow users to freely share, modify, and use this Dataset while maintaining this same freedom for others, provided that the original source and author(s) are credited.
Download file (144.08 kb)
Supplementary material 2 

requency distributions of pored lateral-line scales of species of Pempheris distributed in Japan and Taiwan, with P. flavicycla for comparison

Keita Koeda, Manabu Bessho-Uehara

Data type: xlsx

This dataset is made available under the Open Database License (http://opendatacommons.org/licenses/odbl/1.0/). The Open Database License (ODbL) is a license agreement intended to allow users to freely share, modify, and use this Dataset while maintaining this same freedom for others, provided that the original source and author(s) are credited.
Download file (10.62 kb)
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