Pempheris adusta Bleeker, 1877

Koeda, Keita & Bessho-Uehara, Manabu, 2024, A review of the genus Pempheris (Teleostei, Pempheridae) found in Japan and Taiwan, ZooKeys 1220, pp. 123-163 : 123-163

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. 2013 a: 235; Koeda et al. 2013 b: 221, fig. 1; Koeda et al. 2013 c: 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. 2016 a: 519; Koeda et al. 2016 b: 50, fig. 224; Koeda et al. 2016 c: 8, fig. 3 - G; Koeda and Motomura 2017 a; Koeda 2017 a: 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 2018 a: 193, unnumbered figs; Koeda 2018 b: 298, unnumbered fig.; Koeda 2018 c: 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 2020 a: 407, unnumbered figs; Koeda 2020 b: 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. 2006 a: 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. 2010 a: 75; Motomura et al. 2010 (in part): 131; Koeda et al. 2012 a: 71; Koeda et al. 2012 b: 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. (2013 b). 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 4 ½ – 5 ½ (usually 4 ½); 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.

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.

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. (2013 b). 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 4 ½ (vs usually 5 ½ in Indian Ocean; 4 ½ 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 16 S 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 16 S 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.

Figures

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.

Figure 3. The Maximum-Likelihood (ML) tree of Pempheris species recovered from mitochondria 16 S 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.