Bull. Natl. Mus. Nat. Sci., Ser. A, Suppl. 1, pp. 29–49, March 22, 2007

Review of the Species of the Genus Dentex (Perciformes: Sparidae)
in the Western Pacific Defined as the D. hypselosomus complex
with the Description of a New Species, Dentex abei anda Redescription of Evynnis tumifrons
Yukio Iwatsuki1, Masato Akazaki2 and Nobuhiko Taniguchi3
1

Division of Fisheries Sciences, Faculty of Agriculture, University of Miyazaki,
1–1, Gakuen-kibanadai-nishi, Miyazaki 889–2192, Japan
E-mail: yuk@cc.miyazaki-u.ac.jp
2
Deceased on 12 May 1999
3
Laboratory of Applied Population Genetics, Graduate School of Agriculture, Tohoku University,
Tutumidori, Amamiya-cho, Aoba-ku, Sendai-City, 981–8555 Japan
E-mail: nobuhiko@bios.tohoku.ac.jp

Abstract The lectotype of Chrysophrys tumifrons designated by Boeseman (1947) is a specimenof the species currently recognized as Evynnis japonica Tanaka, 1931. The name is, therefore, thesenior synonym for that species. Dentex spariformis Ogilby, 1910, based on specimens collected at
Moreton Island, Queensland, Australia, is redescribed and resurrected as a valid species, having
previously been synonymized with D. tumifrons (Temminck and Schlegel, 1843). Dentex abei sp.
nov. is described from ten type and 30 non-type specimens collected in the Ryukyu Islands,
Chichi-jima of Ogasawara Islands (Bonin Islands), Japan, and Luzon Island, Philippines. These
two Dentex species, together with D. hypselosomus Bleeker, 1854 and D. fourmanoiri Akazaki and
Séret, 1999 are recognized as a “Dentex hypselosomus complex” and reviewed on the basis ofwestern Pacific and southern Indonesian material. Dentex hypselosomus differs from the others inhaving three eye-sized fluorescent yellow blotches dorsally, plus a small yellow spot posteriorly on
the soft dorsal fin base (vs. small bright cobalt or blue-spotted lines on the body in D. abei and no
spots in D. fourmanoiri and D. spariformis) and deeper suborbital depth. Dentex fourmanoiri differs from D. spariformis in having 13 dorsal fin spines and 9 soft dorsal fin rays (vs. 12 spines and
10 rays). Dentex spariformis has a larger eye than D. abei and D. hypselosomus (mean orbit diameter 2.78 in head length vs. 3.66 and 3.32, respectively). Although none of the four species havebeen recorded in the equatorial zone, D. abei and D. hypselosomus are allopatric in the Northern
Hemisphere, while D. spariformis and D. fourmanoiri are allopatric in the Southern Hemisphere,
too. A key to the complex is provided.
Key words : Taxonomy, Sparidae, Dentex, review, Dentex abei sp. nov.

Akazaki and Séret (1999) considered thesparid genus Dentex to comprise seven Atlantic
and three Pacific species, including their new
species, D. foumanoiri. The Western Pacific Dentex tumifrons (Temminck and Schlegel, 1843) has
long been recorded as a valid species to the East
Asian Shelf (type locality: Nagasaki, Japan). Although Bleeker (1854) described a second, Dentex hypselomus (as hypselosoma), it was considered to be a junior synonym of D. tumifrons by

Akazaki (1962).
Dentex spariformis Ogilby, 1910, describedfrom Moreton Island, Queesland, Australia andrecognized by Fowler (1933), was also synonymized with D. tumifrons by Akazaki (1962).
However, Akazaki (1962) did not comment the
species. Gloerfelt-Tarp and Kailola (1984) identified the species as D. tumifrons and subsequentauthors have followed that view (Allen and
Swainson, 1988; Allen, 1997; Akazaki and Séret,30

Y. Iwatsuki, M. Akazaki and N. Taniguchi

1999; Carpenter, 2001), although Hutchins(2001) transfered it to the monotypic genus Allotaius (Whitley, 1937: type species, Dentex
spariformis) and also considered it to be a valid
species. However, he did not provide an adequate
description and explanation.
Gushiken (1972) reported D. tumifrons fromthe Ryukyu Islands, Japan, but Akazaki (1984)
doubted the identification, referring only to Dentex sp. specimens lacking large yellow blotches
dorsally. Specimens of “D. tumifrons” collectedfrom southern Japan (except the Ryukyu Islands), South Korea, China, and Taiwan clearly
have those markings. Subsequent authors followed Akazaki’s (1984) opinion (Shen, 1993;
Hayashi, 1993, 1999, 2002; Youn, 2002).
Taniguchi et al. (1986) studied genetic differences between the Okinawan and Southern
Japanese (excluding the Ryukyu Islands) populations using allozyme and protein markers, finding a level of difference between them at the subspecies — specific level. However, they did notpropose a new taxon. Recently, Akazaki and
Séret (1999) descibed Dentex fourmanoiri, from
New Caledonia (including Chesterfield Islands),
an endemic species with 13 dorsal fin spines andcharacteristically large eyes, and included an account of Akazaki’s (1984) Dentex sp. of the
Ryukyu Islands, Japan as a separate valid
species. However, there has not been an inclusive
review of the above species, encompassing
known types and non-type specimens throughout
the western Pacific.
In 1947, Boeseman designated a lectotype anda paralectotype from the two syntypes of D. tumifrons. The lectotype has proven to be thespecies currently recognized as Evynnis japonica
Tanaka, 1931. According to Article 74.7.3 of theInternational Commission of Zoological Nomenclature (1999, 2003), D. tumifrons is an available
name and a senior synonym of E. japonica. Accordingly, Dentex hypselosomus Bleeker, 1854 isa valid name. Dentex hypselosomus, D. spariformis, and D. fourmanoiri are herein redescribedas valid species. In addition, a new species ofDentex is described on the basis of specimens

collected from the Ryukyu and Ogasawara Islands (Bonin Islands), Japan. The 4 species together are regarded as the “Dentex hypselosomus
complex.” The unique allopatric distributional
pattern of the 4 species is discussed and a key isprovided. Evynnis tumifrons is redescribed as avalid species.
Materials and Methods
Counts and measurements generally followed
Akazaki (1962) and Akazaki and Séret (1999).
The dorsalmost, spine-like element in the pectoral fin is usually unsegmented but regarded as a
ray in the pectoral fin counts. Standard length
was abbreviated as SL. Institutional codes follow
Leviton et al. (1985) except for the Fisheries Science Course, Faculty of Agriculture, Universityof Miyazaki (MUFS).
Dentex hypselosomus complex
Dentex abei sp. nov., D. fourmanoiri, D.
hypselosomus, and D. spariformis are moreclosely related to each other than to other speciesof Dentex in having the following combination ofcharacters: Dorsal fin rays XII–XIII, 9–10, four
anteriormost dorsal fin spines progressively
longer, third or fourth longest; anal fin rays III, 8,
second spine slightly longer than third; a row of
sharp conical teeth in both jaws, 4 in upper jaw
and usually 6 in lower jaw enlarged, caniform, no
molars; interorbital scales extending almost tolevel with anterior margin of eye; preopercularflange fully scaled; vertebrae 101424; bodyand head red or rosy or pinkish-red with yellow
or reddish-yellow suffusion on dorsal fin rays.
Dentex abei sp. nov.
(Proposed English name: Yellowfin sea brjeam)(Japanese name: Kibire-akarenko)(Figs. 1A, 2A)
Dentex tumifrons (not of Temminck and Schlegel):
Fowler, 1933: 123 (Luzon Island, Philippines);
Gushiken, 1972: 41, fig. 202 (Ryukyu Islands, Japan).
Dentex sp.: Akazaki, 1984: 177, pl. 166B (Okinawa,Dentex in Western Pacific31

Fig. 1. Species of the Dentex hypselosomus complex. A: Dentex abei sp. nov., NSMT-P 57150, holotype,
269 mm SL, Okinawa, Japan; B: D. fourmanoiri, MNHN 1998-301, 226 mm SL, Chesterfield Island, New
Caledonia; C: D. hypselosomus, MUFS 22047, 165 mm SL, Miyazaki, Kyushu Island, Japan; D: D. spariformis, QM 9999, 193 mm SL, off Moreton Bay, Australia.

Japan); Hayashi, 1993: 749 (Ryukyu Islands, Japan);
Akazaki and Séret, 1999: 180 (Ryukyu Islands, Japan);Hayashi, 1999: 859 (Ryukyu and Ogasawara Islands,
Japan); Hayashi, 2002: 859 (Ryukyu Islands and Ogasawara Islands, Japan).
Holotype. NSMT-P 57150, 269 mm SL, male, off
Okinawa Island, Ryukyu Islands, Japan, 50 m depth.
Paratypes. MUFS 7487, 7489, 7491, 7492, 4 specimens, 205–213 mm SL, off Iheiyajima, Ryukyu Island,
Japan; MUFS 17643–17648, 5 specimens, 265–307 mm
SL, off Chichi-jima, Ogasawara (Bonin) Island, Japan, ca.
100 m depth, hand-line.
Non-type specimens. 30 specimens, 120–306 mm.
MUFS 1866, 213 mm SL, Naha fish market, Okinawa,
Japan; MUFS 2341, 182 mm SL, Chi-Lung fish market,
collected off Chi-Lung, northeastern Taiwan, hand-line,
ca. 50 m depth; MUFS 2691, 4524, 4525, 5260, 5262,
5414, 6 specimens, 120–230 mm SL, Naha fish market,
Okinawa; MUFS 17590–17599, 17639–17642, 17649–
17651, 17 specimens, 223–306 mm SL, data as for MUFS
17643–17648; USNM 183185, 201 mm SL, San Fernando,
Union Prov., Luzon Island, Philippines; USNM 191189, 4
specimens, 143–163 mm SL, off I-Lan, northeastern Taiwan.

Diagnosis.

A species of the Dentex hypselo-somus complex with the following combinationof characters: dorsal fin rays XII, 10; pectoral fin
rays 15; orbit diameter small (mean 3.66, range
3.09–4.09 in head length; Fig. 5); snout angle
blunt (range 79°–83°, mean 81°); body prominent pinkish-red with silvery reflection, 5 or 6
bright cobalt or blue small-spotted lines (Fig.
2A) between each horizontal scale row below lateral line, visible only from oblique dorsal view,similar line of small blue spots rising obliquely
above lateral line; eye uniformly reddish-gold
sheen; no prominent yellow blotch in the anterior
nostril and snout region; dorsal fin rays with saffron-yellow membrane but each ray reddish-yellow.
Description. Meristic values and proportional measurements as percentages of standard
length of the holotype, nine paratypes, and 30
non-type specimens of Dentex abei sp. nov. aregiven in Table 1. Characters given in the diagnosis are not repeated.
Body compressed, deep to somewhat slender;

32

Y. Iwatsuki, M. Akazaki and N. Taniguchi
Table 1. Counts and proportional measurements of type and non-type specimens of Dentex abei sp. nov., and D.
fourmanoiri Akazaki and Séret, 1999, expressed as percentages of standard length (means in parentheses).
Dentex abei sp. nov.
Holotype
NSMT-P 57150
n1
Dorsal and anal fin rays
Pelvic and pectoral fin rays
Pored lateral line scales
Scales of transverse series
Scales above and below
lateral line
Gill rakers

Paratypesan9Dentex fourmanoiri
Non-type
specimens
n30

Type
specimensb
n16

Other
specimensn3

XII, 10; III, 8 XII, 10; III, 8 XII, 10; III,
XIII, 9; III, 8 XIII, 9; III, 8
7–8 (7, 1 of 21
specimens)
I, 5; 15I, 5; 15I, 5; 15I, 5; 14–15 I, 5; 15(14, 3 of 16
specimens)
49
48–49
48–50
47–50
49–51
1
1
1
4 /2
4 /2
4 /2–5
41/2–5
41/2–5
6/15
6/14–15
5–6/12–15
5–6/12–15 5–6/14

Squamation on preopercular flangeStandard length (mm)

812
20
Fully scaled269

812–13=20–21
Fully scaled
205–307

7–812–13
19–21
Fully scaled
128–307 (236)4–710–11
14–18Fully scaled
103–226

6–711–12
17–19Fully scaled
202–223

Body depth
Head length
Body width at pectoral fin base
Snout length
Orbit diameter
Dermal eye opening
Suborbital depthInterorbital width
Upper jaw length
Caudal peduncle depth
Caudal peduncle length
Predorsal length
Preanal length
Prepelvic length
Dorsal fin baseAnal fin basePelvic fin spine length
First pelvic fin ray length
Pectoral fin ray length
First dorsal fin spine length
Second dorsal fin spine length
Third dorsal fin spine length
Fourth dorsal fin spine length
First dorsal fin ray length
First anal fin spine length
Second anal fin spine length
Third anal fin spine length
First anal fin ray length

51
36
18
15
12
10
9
11
14
12
16
46
64
40
56
21
16
24
41
7
12
14
14
11
8
13
12
10

42–53
33–36
16–19
12–15
9–10
7–10
7–9
8–11
13–15
10–12
15–19
42–47
69–72
38–42
53–58
18–22
13–17
20–24
33–38
7–8
9–13
13–17
14–18
11–14
6–9
12–14
10–12
10–12

42–53 (46)
33–37 (35)
12–20 (17)
12–16 (14)
9–12 (10)
7–12 ( 9 )
7–9 (8)
9–12 (10)
12–16 (14)
10–13 (12)15–20 (18)
42–55 (46)64–73 (70)38–45 (40)
51–58 (54)
18–23 (20)
13–19 (15)
20–25 (22)31–41 (37)
4–8 ( 7 )8–14 (11)
13–19 (15)
13–21 (16)
11–14 (12)
6–10 ( 8 )11–16 (13)
10–14 (11)
9–13 (11)

44–49
34–38
13–19
—
—
—
—
—
—
—
—
—
—
—
—
—
—
—
34–41
—
—
—

44–45
35–37
16–19
13–15
11–13
11–12
7.6–8.1
10–11
15–16
12–13
19–20
42–46
64–71
39–45
51–53
20
16–17
20–21
36–37
7–9
10
14–15
15–16
12
8–9
14–15
12–13
12

a

—
—
—
—

MUFS 7487, 7489, 7491–7492, 4 specimens, 205–213 mm SL, off Iheiyajima, Ryukyu Is., Japan; MUFS
17643–17648, 5 specimens, 265–307 mm SL, off Chichi I., Ogasawara (Bonin) Is., Japan; b Data based on Akazaki and
Seret (1999).

Dentex in Western Pacificanterodorsal profile ascending sharply, rostro-occipital profile somewhat convex; orbit small diameter, slightly greater than interorbital width;
snout length much greater than orbit diameter;maxilla reaching beyond vertical through anterior
margin of eye; mandibular profile almost
straight; preopercle entire with rounded angle;
preopercular flange fully scaled; first to third dorsal fin spines progressively longer, third or fourth
dorsal fin spine longest; predorsal length clearlyless than body depth, slightly less than dorsal finbase length; pectoral fin tip reaching to or slightly beyond first anal fin spine base, its length
greater than head length; anal fin spines stout,
second longest, longer than orbit diameter; caudal fin weakly forked, lobes short, broad; mostupper gill rakers on lower limb of gill arch long,a few lower gill rakers sometimes shortened or
rudimentary.
Color when fresh. Based on color transparencies and observations of type and non-type specimens, in addition to description presented inTable 4: pectoral fin rays strongly pinkish-red onmuch of upper part of fin; anal fin rays uniformly
reddish-yellow (anteriorly and along posterior
basal portion), remainder hyaline; caudal fin rays
mostly prominent pinkish-red or yellow.
Color of preserved specimens. Upper sidesof head and trunk uniformly yellowish-tan; darkbrownish small-spotted lines between each horizontal scale row below lateral line (not discernible in some long term preserved specimens).
Distribution. Dentex abei sp. nov. is known
only from the Ryukyu Islands, including the Okinawa Trough, Ogasawara Islands, Japan, northeastern Taiwan, and off Luzon Island, Philippines(Fig. 6). Attempts to find specimens from otherlocalities were unsuccessful. Most examples ofD. abei were collected by hand-line in depths of
50–150 m.
Etymology. The specific name “abei” recognizes the late Dr. Tokiharu Abe, who studied
many Japanese fish species and was aware of theuniqueness of this species.
Remarks. On the basis of isozyme and pro-

33

Fig. 2. Body coloration. A: Dentex abei, holotype,
MSMT-P 57150, 269 mm SL; B: D. hypselosomus, MUFS 22984, 184 mm SL.

tein markers, Taniguchi et al. (1986) concluded
that “The genetic distances between Dentex tumifrons [hypselosomus] and Dentex sp. [D.
abei] were larger than the subpopulation level but
smaller than the species level” but did not propose a new name for the latter. The recognitionof D. abei as a valid species is based on meristic
and morphological information, as well as color
characters (see Discussion below and Remarks
for D. hypselosomus).
The specimen from the Philippines reported by
Fowler (1933), Dentex hypselosomus (as tumifrons; USNM 183185, 201 mm SL, San Fernando, Union Prov., Luzon Island, Philippines) is
D. abei.
Dentex fourmanoiri Akazaki and Séret, 1999
(English name: Fourmanoir’s sea bream)(Fig. 1B)

Dentex fourmanoiri Akazaki and Séret, 1999:
179, figs. 1–2 (type locality: off East of Chesterfield Island, New Caledonia, 19°27S, 158°34E,

34

Y. Iwatsuki, M. Akazaki and N. Taniguchi

217 m depth); Carpenter, 2001: 2999 (New Caledonia including Chesterfield Island).
Types. Type specimens were not examined
but raw data for them were provided by the late
M. Akazaki, prior to his death.
Non-type specimens. MUFS 6032, 6084,
6085, 3 specimens, 199–228 mm SL, off New
Caledonia.
Diagnosis. A species of the Dentex hypselosomus complex with the following combinationof characters: dorsal fin rays XIII, 9; pectoral fin
rays 14–15; orbit diameter large (range 2.40–3.35
in head length, mean 2.78; Fig. 4); snout, sharp
angle (range 70°–83°, mean 77°); body rosy topale red with silvery reflection, often with 7 or 8
wide purplish-silver longitudinal stripes between
adjacent horizontal scales below lateral line
(small blue-spotted lines absent), visible fromlateral view; eye silvery, some specimens withyellowish tint ventrally without a prominent yellow blotch in anterior nostril and snout region;
dorsal fin rays pale yellowish with bright yellow
distal margin.
Description. Meristic values and proportional measurements as percentage of standard length
for the holotype, paratypes, and 3 non-type specimens of Dentex fourmanoiri are given in Table
1. Characters presented in the diagnosis are notrepeated.
Body compressed, deep to somewhat slender;
anterodorsal profile ascending gradually, rostrooccipital profile somewhat convex, body somewhat thickened; orbit diameter large, slightly
greater than interorbital width; snout length
much greater than orbit diameter; maxilla reaching to or slightly beyond vertical through anterior
margin of eye; mandibular profile almost
straight; preopercle entire with rounded angle;
preopercular flange fully scaled; first to third dorsal fin spines becoming progressively longer,
fourth dorsal fin spine longest; predorsal length
less than half body depth, clearly less than dorsal
fin base length; pectoral fin tip reaching to orslightly beyond first anal fin spine base, its length
subequal to head length; anal fin spines robust,second longest, longer than orbit diameter; cau-dal fin weakly forked, lobes short, broad; mostupper gill rakers on lower limb of gill arch long,a few lower gill rakers sometimes abruptly shortened or rudimentary.
Color when fresh. Based on color transparencies, reproduced as, Fig. 1B and fig. 2 of
Akazaki and Séret, 1999 of type and non-type
specimens: a render part of pectoral and anal fin
rays uniformly pale, caudal fin rays pale yellowish with bright yellow distal margin.
Color of preserved specimens. Upper sidesof head and trunk uniformly yellowish-tan; mostlife colors not discernible but 7 or 8 wide silver(guanine) longitudinal stripes between adjacenthorizontal scale rows below lateral line in some
ethyl-alcohol specimens.
Distribution. The holotype of Dentex fourmanoiri was collected at a depth of 217 m, Eastof Chesterfield Island, New Caledonia with otherspecimens coming from the main islands of New
Caledonia (Fig. 6). The species has not been reported elsewhere.

Dentex hypselosomus Bleeker, 1854
(English name: Yellowback sea-bream)(Japanese name: Kidai)(Figs 1C, 2B, 3A)
Chrysophrys tumifrons Temminck and Schlegel, 1843: 70,
pl. 34 [Nagasaki Bay, Japan; in part, only for paralectotype of Boeseman (1947) but justified as unavailable
name in this species because of lectotype of Boeseman(1947) identified as present Evynnis japonica Tanaka,
1913, this species being a senior synonym of Evynnis
tumifrons in this study; see Remarks].
Dentex hypselosoma Bleeker, 1854: 402 (type locality:
Nagasaki, Japan); Steindachner, 1896: 199 (Kobe,
Hiogo, Nagasaki).
Synagris hypselosoma: Bleeker, 1873: 40 (Nagasaki,
Japan); Bleeker, 1875–1876: 314, fig. 2, pl. 36 (Nagasaki, Japan).
Pagrus tumifrons: Ishikawa and Matsuura, 1897: 53(Tokyo); Jordan and Snyder, 1901: 79 (Nagasaki).
Dentex hypselosomus: Jordan and Evermann, 1902: 345,
fig. 17 (Taiwan); Jordan and Richardson, 1909: 185(Takao).
Taius tumifrons; Jordan and Thompson, 1912: 531(Japan); Snyder, 1912: 415 (Kagoshima, Japan); Jordan and Metz, 1913: 34 (Tsushima Straits); Tanaka,Dentex in Western Pacific35

Fig. 3. Lectotypes of Dentex hypselosomus, RMNH 5678, 138 mm SL (A) and Dentex spariformis, QM I. 1549,
172 mm SL (B).
1915: 374, pl. 102, fig. 314 (southen Japan, Taiwan,
China); Jordan and Hubbs, 1925: 241 (Tokyo, Kobe,
Kagoshima. Misaki. Toyama, Miyazu); Oshima, 1927:
131, figs. 2–3 (Keelung, Tong King Bay); Fowler,
1930: 592 (Nagasaki, Japan), 596 (Shanghai, China),
609 (Hong Kong, China).
Raius tumifrons: Jordan and Thompson, 1914: 256 (Misaki, Tsushima Straits).
Dentex tumifrons: Akazaki, 1962: 137 (southern Japan);
Akazaki, 1984: 177 (southern Japan); Okamura et al.,
1985: 511, 689 (Okinawa Trough, Japan); Chen et al.,
1997: 92 (Nansha Islands to South China); Akazaki

and Séret, 1999: 177 (southern Japan); Carpenter,
2001: 619 (South China Sea); Sadovy and Cornish,
2000: 137 (Hong Kong, China); Shinohara et al., 2001:
326 (Tosa Bay, Japan); Carpenter, 2001: 3000 (Japan,
in part); Hayashi, 2002: 859 (southern Japan); Youn,
2002: 342, 645 (South Korea).
Lectotype. RMNH 5678 (Fig. 3A), 138 mm SL, Nagasaki, Japan.
Paralectotype. RMNH 35385, 110 mm SL, Nagasaki, Japan.
Non-type specimens. 51 specimens, 78–306 mm SL.
MUFS 1443, 112 mm SL, off Miyazaki City, Miyazaki,

36

Y. Iwatsuki, M. Akazaki and N. Taniguchi

Kyushu Island, Japan; MUFS 2342, 212 mm SL, Chi-Lun,
Taiwan; MUFS 3972, 78 mm SL, Shirahama, Wakayama,
Honshu Island, Japan; MUFS 4322, 154 mm SL, Kushima, Miyazaki, Kyushu Island, Japan; MUFS 4519–4521,
3 specimens, 148–162 mm SL, Kawaminami, Miyazaki,
Japan; MUFS, 5263, 5265–5272, 5275–5276, 11 specimens, 120–150 mm SL, Ushibuka, Kumamoto, Kyushu
Island, Japan; MUFS 12086–12088, 3 specimens, 97–112mm SL, Nishimaizuru, Kyoto, Honshu Island, Sea ofJapan; MUFS 17651–17669, 18 specimens, Meitsu,
Miyazaki, Kyushu Island, Japan; MUFS 12751–12752, 2
specimens, 124–127 mm SL, southern Kagoshima Bay,
Kagoshima, Kyushu Island, Japan; MUFS 19870, 190 mm
SL, off Miyotoura, Nichinan, Miyazaki, Kyushu Island,
Japan; MUFS 17996, 270 mm SL, off Oshima, Nango,
Miyazaki, Japan; MUFS 22984, 184 mm SL, off Oshima,
Nango Miyazaki, Kyushu Island, Japan; RMNH 960(paralectotype of Boeseman, 1947), 141 mm SL, Nagasaki Bay, Japan; SMF 25471, 143 mm SL, off Hainan Island, China, South China Sea.

Diagnosis. A species of the Dentex hypselosomus complex with the following combinationof characters: dorsal fin rays XII, 10; pectoral fin
rays usually 15 (rarely 16); orbit diameter small,range 2.66–3.87, mean 3.32, in head length (Fig.
5); snout angle blunt, range 82°–88°, mean 83°;
body red dorsally, becoming pale red ventrallywith a yellowish gold sheen, snout vivid yellow,
three large fluorescent yellow blotches dorsally
beneath dorsal fin base and a single black irissized yellow spot on base of posteriormost soft
dorsal fin ray (Figs. 1C, 2B), clearly visible fromoblique dorsal view (Fig. 2B); eye uniformly
prominent reddish-gold; prominent yellow blotch
on anterior nostril and snout region; dorsal finmembrane and rays orange-yellow to pale red butwith a longitudinal hyaline stripe beneath finbase, and soft membrane and rays saffron-yellow.
Description. Meristic values and proportional measurements as percentages of standard
length for the lectotype and non-type specimensof Dentex hypselosomus are given in Table 2.
Characters presented in the diagnosis are not repeated.
Body compressed, deep to somewhat slender;
anterodorsal profile ascending sharply, rostro-occipital profile somewhat convex; orbit diametersmall, subequal to or slightly less than interorbital width; snout length much greater than orbit

diameter; maxilla reaching beyond verticalthrough anterior margin of eye to verticalthrough anterior margin of iris; mandibular profile almost straight; preopercle entire with rounded angle; preopercular flange fully scaled; first to
third dorsal fin spines progressively longer, thirdor fourth dorsal fin spine longest; predorsal
length less than body depth, less than dorsal finbase length; pectoral fin tip reaching to or slightly beyond first anal fin spine base, its length
greater than head length; anal fin spines stout,
second longest, longer than orbit diameter; caudal fin weakly forked, lobes short, broad; mostupper gill rakers on lower limb of gill arch long,a few lower gill rakers sometimes abruptly shortened or rudimentary.
Color when fresh. Based on color transparencies of many specimens, including MUFS
22047 (Fig. 1C), 165 mm SL, and observationsof many non-type specimens, in addition to description presented in Table 4: pectoral fin rays
orange-yellow to reddish for much of upper partof fin; anal fin rays reddish-yellow anteriorly, orange-yellow or yellow posteriorly, except forhyaline rays and lower distal margin hyaline; caudal fin rays mostly red to pale red, yellowish anteriorly.
Color of preserved specimens. Upper partsof head and trunk uniformly yellowish-tan; life
colors lost.
Distribution. Dentex
hypselosomus
isknown from southern Japan (except the Ryukyu
and Ogasawara Islands), southern Korea, China,
and western Taiwan (Fig. 6). Attempts to locate
specimens from the Indian Ocean and southeastern Asia to Australia were unsuccessful. The
species is highly esteemed commercially, and it
is caught mainly between 50 and 200 m in the
East China Sea, where it has two spawning seasons per year in June and November (Yamada,1986).
Remarks. Chrysophrys tumifrons was described on the basis of 2 syntypes (RMNH D418,
stuffed, 250 mm SL and RMNH 960, 141 mm
SL) by Temminck and Schlegel (1843). Subsequently, Boeseman (1947) designated a lectotype

Body depth
Head length
Body width at pectoral fin base
Snout length
Orbit diameter
Suborbital depth
Dermal eye opening
Interorbital width
Upper jaw length
Caudal peduncle depth
Caudal peduncle length
Predorsal length
Preanal length
Prepelvic length
Dorsal fin baseStandard length (mm)Pored lateral line scales
Scales above and below lateral line
Scales between 5th dorsal fin
spine base and lateral line
Squamation on preopercular flange
Gill rakers

Pelvic and pectoral fin rays

Dorsal and anal fin rays

48
34–35
16–17
13
11–12
8–10
12–13
9–11
13–14
12
20
45
61
39–40
52

Fully scaled
7–81320–21
133–144

Fully scaled
713
20
172

47
35
15
12
10
8
11
10
13
12
19
44
64
41
50

48–49
7/15
41/2

49
7/15
41/2

XII, 10;
III, 8I, 5; 15n2

n1

XII, 10;
III, 8I, 5; 15

Paralectotypes
QM I.477 andAMS I.12538

Lectotype
QM I. 1549

Dentex spariformis

42–54 (48)
34–39 (36)
14–28 (17)
10–16 (14)
10–15 (13)
7–10 (8)
9–14 (12)
8–13 (11)
12–17 (14)
10–13 (12)
14–24 (20)42–52 (45)61–75 (66)
37–48 (42)48–56 (52)

Fully scaled
6–810–13
16–21
61–283 (131)

XII, 10;
III, 8I, 5; 15–16(16, 11 of 22
specimens)
47–49
5–6/13–16
41/2

n39

Non-type
specimens

Lectotype andparalectotype of
Synagris
hypselosoma
RMNH 5678
n2

52
34
16
14
14
11
8
—
11
9
11
7
12
16
16

Fully scaled
712
19
141

46–48
35
13
13
12–13
9–10
11–12
8
13–14
12–13
20–22
45
63–64
39
53–54

Fully scaled
712–1319–20
110–138

XII, 10;
XII, 10;
III, 8
III, 8I, 5 and III, 5 inI, 5; 15 or 16
right (as deformity);
15
48
48 or 49
5/15
5–6/15–16
41/2
41/2

n1

Lectotype ofDentex tumifrons
RMNH 960an51

Non-type
specimens

41–53 (48)
29–35 (35)
14–20 (17)
11–18 (15)
9–14 (11)
7–11 (8)
8–11 (10)
9–12 (10)
10–17 (14)11–20 (12)16–22 (19)43–49 (46)58–74 (64)37–45 (39)49–59 (53)Fully scaled
6–910–12
16–21
78–70 (169)47–49
5/13–15
41/2

XII, 9–10 (9, 1 of
50 specimens); III, 8
I, 5; 15–16(16, 4 of 50 specimens)

Dentex tumifrons

Table 2. Counts and proportional measurements of type and non-type specimens of Dentex spariformis Ogilby, 1910 and D. tumifrons (Temminck and Schlegel,1843), expressed as percentages of standard length (means in parentheses).

Dentex in Western Pacific
37

a

20
18
24–25
36–37
8
12
16–17
13
8–9
14–15
13–14
14

7
13
12
11
—

n1

19
15
24
36
7
13
14

n2

Lectotype
QM I. 1549

Boeseman’s (1947) paralectotype (see text).

Anal fin basePelvic fin spine length
First pelvic fin ray length
Pectoral fin ray length
First dorsal fin spine length
Second dorsal fin spine length
Third dorsal fin spine length
Fourth dorsal fin spine length
First dorsal fin ray length
First anal fin spine length
Second anal fin spine length
Third anal fin spine length
First anal fin ray length

Paralectotypes
QM I.477 andAMS I.12538

Dentex spariformisn39

Non-type
specimens(Continued).

11–15 (13)
7–10 ( 8 )
12–17 (14)
11–16 (13)
9–16 (14)

18–24 (20)
14–20 (17)20–28 (24)
25–42 (37)
6–9 2( 8 )8–15 (11)
13–18 (16)Table 2.

14
13
13
12
16
25
14
16
47
62
—
—
—

n1

Lectotype ofDentex tumifrons
RMNH 960a

18–20
15–17
20
39
8–9
14
17–18
16–18
13
8–9
13–14
13
12

Lectotype andparalectotype of
Synagris
hypselosoma
RMNH 5678
n2

n51

Non-type
specimens

18–24 (20)
14–18 (17)19–25 (22)34–43 (38)
5–11 ( 8 )8–14 (12)13–18 (16)
15–18 (17)
11–15 (13)
4–10 ( 8 )10–15 (12)
9–15 (11)
10–14 (11)Dentex tumifrons

38
Y. Iwatsuki, M. Akazaki and N. Taniguchi

Dentex in Western Pacific(Fig. 4A) for the larger specimen. It has provento be an example of a different genus andspecies, long recognized as Evynnis japonica
Tanaka, 1931, as well as by Yamaguchi and
Machida (2003). The specimen has 61 pored lateral line scales (clearly countable although enameled) (vs. 48–50 in D. hypselosomus), 9 soft anal
fin rays (vs. 8 in D. hypselosomus), lacks scales
on the preopercular flange (vs. fully scaled in D.
hypselosomus), and the first and second dorsal fin
spines considerably shorter than the third andfourth dorsal fin spines, all characteristics ofEvynnis japonica (Tables 2–3; Tanaka, 1931;
Akazaki, 1962; Carpenter, 2001).
The smaller paralectotype (RMNH 960, 141
mm SL) is a specimen of Dentex hypselosomus
as recognized in this study (Table 2), with characteristics matching those presented by Akazaki(1962) and Carpenter (2001). Evynnis japonica
is, therefore, a junior synonym of Evynnis tumifrons (Table 3). Dentex hypselosomus Bleeker,
1854 based on 2 syntypes (RMNH 5678,
110–138 mm SL) from Nagasaki, Japan is the
only other available name for this species. Bleeker’s description of D. hypselosomus clearlymatches (Table 2). We herein designate the larger
syntype (RMNH 5678, 138 mm SL) of Dentex
hypselosomus (originally as hypselosoma) as thelectotype, the smaller specimen becoming theparalectotype (RMNH 35385, 110 mm SL).
Dentex spariformis Ogilby, 1910
(Proposed English name: Saffronfin sea bream)(Fig. 1D)
Dentex spariformis Ogilby, 1910: 91 [ca. 66.7 km (as 36
miles) south 12°W from Moreton Island, Queesland,
Australia, ca. 14 m (7 fathoms)]; Ogilby, 1916: 169, pl.
21 (off Cape Moreton); Fowler, 1933: 124 (Queensland, Australia).
Dentex tumifrons (not of Temminck and Schlegel): Gloerfelt-Tarp and Kailola, 1984: 207, unnumbered color
plate on p. 206 (off Lombok Island, Indonesia); Allen,
1997: 132 (Western Australia); Carpenter, 2001: 3000
(Western Pacific, in part).
Allotaius spariformis: Hutchins, 2001: 35 (Australia).
Lectotype. QM I. 1549, 172 mm SL, ca. 66.7 km (as39

36 miles), south 12°W. from Moreton Island, Queensland(Qld), Australia, ca. 14 m (7 fathoms).
Paralectotypes. AMS I.12538, 143 mm SL, same locality as lectotype; QM I. 477, 133 mm SL, same locality
as the lectotype.
Non-type specimens. 61–283 mm SL, n39. CSIRO
CA 401, 88 mm SL, Cape Lambert, East Australia;
CSIRO CA 3029, 200 mm SL, East Australia; CSIRO H
2017-01, 144 mm SL, Geraldton, Australia; CSIRO H
3644-01, 151 mm SL, Qld, Australia; CSIRO H 4649-20,
160 mm SL, East Australia; FRLM 14692, 246 mm SL,
Bali, Indonesia; NTM 10752-008, 101 mm SL, off southern Lombok Island, Indonesia, 150 m depth, trawl; NTM
11748-003, 100 mm SL, East of Dunk Island, Qld,
Austalia, 218–220 m depth, trawl; NTM 11749-004, 88mm SL, East of Dunk Island, Qld, Australia, 220 m depth,
trawl; NTM S. 11751-010, 151 mm SL, East of Dunk Island, Qld, Australia, trawl, 264 m depth; NTM 11752-012,
3 specimens, 141–169 mm SL, East of Dunk Island, Qld,
Australia, 260–262 m depth, trawl; NTM 12187-003, 269
mm SL, Northwest of Melville Island, NT, Australia,
trawl, 113 m depth; NTM S. 13301-004, 214 mm SL,
western part of Lyndoch Bank, Arafura Sea, Indonesia,
159 m depth, trawl; NTM S. 13660-002, 283 mm SL, Lyndoch Bank, Arafra Sea, Indonesia, 183 m depth, trawl;
NTM 61225-002, 237 mm SL, Australia; QM 12989, 7
specimens, 61–74 mm SL, ca. 13 km (7 miles), off northwest of Cape Moreton, Qld, Australia, 60 fms (ca. 100 m)
depth, trawl; QM I. 18697, 121 mm SL, off eastern Fraser
Island (25°27–17N, 153°46–43E), Qld, Australia,
trawl, 230–183 m depth; QM 19209-001, 2 specimens,
141–152 mm SL, off Sandy Cape (23°59–23°54N,
152°47–152°43E), Qld, Australia; QM I. 25297,
154 mm SL, off Swain Reef (21°22N, 153°05E), Qld,
Australia, trawl; QM 25689, 10 specimens, 61–121 mm
SL, off Swain Reef (21°42N, 152°55E), Qld, Australia,
185–190 m depth, trawl; QM 26532, 225 mm SL, Australia.

Diagnosis. A species of the Dentex hypselosomus complex with the following combinationof characters: dorsal fin rays XII, 10; pectoral fin
rays 15 or 16; orbit diameter large (mean 2.78,
range 2.37–3.39 in head length; Fig. 5); snout
angle sharp (mean 76°, range 69°–84°); bodycolor light pinkish-red with silvery reflection, no
obvious blue-spotted lines or yellow blotches (asin D. abei) or wide longitudinal purplish-silver
stripes between each horizontal scale row below
lateral line (as in D. fourmanoiri); eye color generally pinkish-gold or with silver sheen (lower
part yellowish in some specimens); prominent

40

Y. Iwatsuki, M. Akazaki and N. Taniguchi

yellow blotch absent from anterior nostril andsnout region; dorsal fin rays saffron-yellow onmembrane and rays.
Description. Meristic value and proportional
measurements as percentages of standard length
for the holotype, two paratypes, and 39 non-type
specimens of Dentex spariformis are given in
Table 2. Characters presented in the diagnosis are
not repeated.
Body compressed, deep to somewhat slender;
anterodorsal profile ascending gradually, rostrooccipital profile somewhat convex, body somewhat thickened; orbit diameter large, slightly
greater than interorbital width; snout length
much greater than orbit diameter; maxillary
reaching through vertical level of anterior marginof eye; mandibular profile almost straight; preopercle entire with rounded angle; preopercularflange fully scaled; first to third dorsal fin spines
becoming progressively longer, third or fourth
dorsal fin spine longest; predorsal length clearlyless than body depth, slightly less than dorsal finbase length; pectoral fin tip reaching to or slightly beyond first anal fin spine base, its length
greater than head length; anal fin spines stout,
second longest, longer than orbit diameter; caudal fin weakly forked, lobes short, broad; mostupper gill rakers on lower limb of gill arch long,a few lower gill rakers sometimes abruptly shortened or rudimentary.
Color when fresh. Based on color transparencies from FishBase as D. tumifrons andphotograph of J. Johnson, in addition to description presented in Table 4: a render (upper) part ofpectoral fin rays saffron-yellow or pale yellow;
anal fin membrane and rays pale saffron-yellow;
caudal fin membrane and rays mostly saffron topale yellow.
Color of preserved specimens. Upper sidesof head and trunk uniformly yellowish-tan; life
colors otherwise lost.
Distribution. Dentex spariformis is currently known only from northern Australia, Bali, andLombok Island and the Arafura Sea, Indonesia(Fig. 6), at depths between ca. 100 and 264 m.
Remarks. Dentex spariformis Ogilby, 1910

has long been considered a junior synonym of D.
tumifrons (D. hypselsomus of this study; e.g.,
Gloerfelt-Tarp and Kailola, 1984; Allen, 1997;
Akazaki and Séret, 1999; Carpenter, 2001) but isherein considered as valid. The species lacks thefollowing characters that are unique to D.
hypselosomus: 3 large fluorescent yellow blotches dorsally below the dorsal fin base plus a blackiris-sized spot on the base of posteriormost soft
dorsal fin ray (Figs. 1C, 1D, 2B).
The largest syntype of Dentex spariformis
Ogilby, 1910 (QM I. 1549, 172 mm SL) is here
designated the lectotype, QM I.477 and AMS
I.12538 (133–144 mm SL) becoming paralectotypes.
Discussion
Comparison. Selected counts and charactersof Dentex abei sp. nov., D. fourmanoiri, D.
hypselosomus, and D. spariformis are comparedin Tables 1 and 2. Dentex fourmanoiri clearly differs from the other 3 species in having 13 dorsal
fin spines and 9 soft dorsal fin rays (vs. 12 dorsal
fin spines and 10 dorsal fin soft rays). Dentex
hypselosomus is similar to D. abei in having asmaller orbit diameter (means 3.32 and 3.36 inhead length, respectively vs. mean 2.78 in D.
fourmanoiri and D. spariformis; Fig. 5A), butdiffers from it in having a greater suborbital
depth (mean 9.5% SL vs. mean 7.8% SL in D.
abei and D. fourmanoiri, 8.1% SL in D. spariformis, see Table 4 and Fig. 5B), 3 large fluorescent yellow blotches dorsally beneath the dorsal
fin ray base, an iris sized yellow spot on aroundthe last soft dorsal fin base (Figs. 1C, 2B), that isclearly visible at an oblique dorsal view (Fig.
2B), and a prominent yellow blotch on the anterior nostril and snout region (vs. yellow blotchesabsent in others). Dentex abei sp. nov. is distinguishable from the three other species in having
5 or 6 bright cobalt or blue small-spotted lines(Fig. 2A) between adjacent horizontal scale rows
below the lateral line that are visible only froman oblique dorsal view, and such lines risingobliquely above the lateral line (vs. small blue-

Dentex in Western Pacificspotted lines absent)(Fig. 2A).
Distribution. Dentex abei sp. nov., D. fourmanoiri, D. hypselosomus, and D. spariformis
have allopatric and non-equatorial distributionsin the western Pacific (Fig. 6). In the Northern
Hemisphere, Dentex abei is currently known only
from the Ryukyu Islands including the Okinawa
Trough, Ogasawara Islands, Japan and northeastern Taiwan, plus off Luzon Island, Philippines,
while D. hypselosomus is currently known fromthe East Asian Shelf, including southern Japan(except the Ryukyu and the Ogasawara Islands),
southern Korea, Taiwan (generally the westernpart of Taiwan), and China. In the Southern
Hemisphere, D. spariformis is distributed off
Australia (except southwestern part), while D.
formanoiri is currently known only from off New
Caledonia, including the Chesterfield Island. A
similar distributional pattern is known for congereric trichiurids (Burhanuddin et al., 2002;
Burhanuddin and Iwatsuki, 2003; Chakraborty etal., 2005) with the Northern Hemisphere: Trichiurus russelli in the northern Gulf of Thailand andnorthern Bengal Bay and T. brevis in southern
China and in the Southern Hemisphere: T.
nikolensis in northwestern Australia, and T. australis in northeastern Australia. Species in both
Dentex and Trichiurus are known to be basicallybenthic and deep-sea species strongly connectedwith a continental shelf (Nakamura and Parin,
1993 for Trichiurus) and are not strong swimmers, like species of the carangid genus Seriola.
In the Northern and Southern Hemisphere, Dentex species have a clearly allopatric distribution
in two small regions, the East Asian Shelf (D.
abei and D. hypselosomus), and northern Australia and New Caledonia (D. fourmanoiri and D.
spariformis).
Generic position. Akazaki (1962) provided
the following combination of characters as a diagnosis for the genus Dentex: a row of sharp conical teeth in both jaws, 4 in upper jaw and 6 inlower jaw enlarged, caniform; no molars;
i445nterorbital scales almost extending to levelwith anterior margin of eye; preopercular flange
fully scaled; a transverse series of scale rows be-

41

tween lateral line and median dorsal spines
41/2–71/2.
Prior to these studies, Jordan and Thompson(1912) and Whitley (1937) each proposed a newgeneric name, Taius (type species: Chrysophrys
tumifrons Temminck and Schlegel, 1843) and Allotaius (type species: Dentex spariformis Ogilby,1910), respectively. Akazaki (1962) synonymized
Polysteganus of Klunzinger (1870, type species:
Polysteganus coeruleopunctatus Klunzinger,
1870), Taius and Opsodentex of Fowler (1925,
type species: Sparus macrophthalmus Bloch,1791) with Dentex (Cuvier, 1814, type species:
Sparus dentex Linnaeus, 1758), recognizing thefirst and last as subgenera. Although interrlationships of all species within the genus remain unresolved, we follow Akazaki (1962) and Carpenter(2001) in regarding D. fourmanoiri and D.
hypselosomus plus D. spariformis and D. abei sp.
nov. considered in this study as a natural group.
Further work is required to understand interrelationships within the Sparidae.
Evynnis tumifrons (Temminck andSchlegel, 1843)(English name: Crimson seabream)(Japanese name: Chidai)(Figs. 4A–C)
Chrysophrys cardinalis (not of Lacepède, 1802): Temminck and Schlegel, 1843: 69, pl. 33 (Nagasaki, Japan;
pl. 33 as Fig. 4B).
Chrysophrys tumifrons Temminck and Schlegel, 1843: 70,
pl. 34 [Nagasaki Bay, Japan; in part, only for lectotypeof Boeseman (1947) but justified as available validname in the genus Evynnis].
Evynnis cardinalis: Jordan and Thompson, 1912: 573(Tokyo, Japan).
Chrysophrys cardinalis (not of Lacepéde, 1802): Tanaka,
1915: 313, pl. 51, fig. 3 (Japan and China).
Evynnis japonica Tanaka, 1931: 29 (no type known, newname for Chrysophrys cardinalis of Temminck andSchlegel, 1843, not of Sparus cardinalis Lacepède,1802); Akazaki, 1962: 176 (Japan); Akazaki, 1984:
177 (Japan, China, Korea, and Taiwan); Hayashi, 1999,
2002: 858; Carpenter, 2001: 3001 (Temperate West Pacific from Hokkaido to East China Sea; a single record
from Bali); Youn, 2002: 342, 616 (Korea).
Lectotype. RMNH D418, stuffed, 250 mm SL,

42

Y. Iwatsuki, M. Akazaki and N. Taniguchi

Fig. 4. Lectotype (A) of Chrysophrys tumifrons, a plate (B) of Chrysophrys cardinalis of Temminck andSchlegel, 1843, and the lectotype (C) of Evynnis japonica. A, RMNH D418 [Boeseman’s (1947) lectotype ofChrysophrys tumifrons Temminck and Schlegel, 1843], stuffed, 250 mm SL, Nagasaki; B, plate 33 of Temminck and Schlegel, 1843; C, RMNH 954b, lectotype, 128 mm SL, Nagasaki, Japan.

Dentex in Western PacificFig. 5. Suborbital depth (A) and orbit diameter(B) relative to standard length in the four
species of the Dentex hypselosomus complex.

Nagasaki, Japan.
Non-type specimens. 126–270 mm SL, all from Japan,n19. MUFS 915, 270 mm SL, Tanabe, Wakayama;
MUFS 11723, 11725, 11727, 11738–11739, 11743,
11745, 12054, 12252–12254, 12569, 12 specimens,
126–240 mm SL, Meitsu, Nango, Miyazaki, Kyushu Island; RMNH D401 (specimen of Temminck andSchlegel’s Chrysophrys cardinalis, and paralectotype of
Evynnis japonica), stuffed, 209 mm SL, Nagasaki, Japan,
coll. Bürger; RMNH D402, stuffed, 253 mm SL, same as
RMNH D401; RMNH D403, stuffed, 204 mm SL, sameas RMNH D401; RMNH 954a, b, c, d (Boeseman, 1947:47), 4 specimens [RMNH 954b, 128 mm SL, herein designated as lectotype of Evynnis japonica; RMNH 954d
(63 mm SL), RMNH 954c (72 mm SL), RMNH 954a (143mm SL)], Nagasaki, Japan, coll. Von Siebold.

Diagnosis. A species of Evynnis with the following combination of characters: dorsal fin rays
XII, 10; anal fin rays III, 9; pectoral fin rays usually 15; first and second dorsal fin spines considerably shorter than third and fourth dorsal fin
spines: third and fourth dorsal fin spines extend-

43

Fig. 6. Distribution of the four species of the“Dentex hypselosomus complex”: Dentex abei
sp. nov. (square), Dentex fourmanoiri (star), D.
spariformis (circle), and D. hypselosomus triangle). Solid and open symbols represent specimens examined and literature records, respectively.

ed but third dorsal fin spine less than 2 time in
fifth dorsal fin spine; no scales on preopercularflange; body depth relatively lower, standard
length/body depth over 2.1; scattered cobalt blue
spotted markings on side in fresh; teeth on vomer
usually present.
Description. Further work is required for
detailed description with a review of the genus
Evynnis including generic definition. We presently put diagnostic characters above in this species
as well as Comparison below.
Distribution. Evynnis tumifrons is currently
known from Hokkaido, Honshu, Shikoku, andKyushu islands, Japan except the Ogasawara and
Ryukyu Islands, southern Korea, Taaiwan, andChina (Akazaki, 1962, 1984; Youn, 2002; Carpenter, 2001). Carpenter (2001) noted a single
record from Vali, Indonesia.
Comparison. In the Indo-Pacific, Evynnis
tumifrons is similar to E. cardinalis Lacepède,

44

Y. Iwatsuki, M. Akazaki and N. Taniguchi
Table 3. Counts, selected characters, and proportional measurements of Evynnis japonica Tanaka, 1913, expressed as percentages of standard length (means in parentheses).
Evynnis japonica
RMNH D918
n1
Dorsal and anal fin rays
Pelvic and pectoral fin rays
Pored lateral line scales

Non-type specimens
in MUFS collection(126–270 mm SL)
n12

XII, 10; III, 9
I, 5; 1561, clearly countable althoughspecimen dried—

XII, 10; III, 9
I, 5; 15
58–61

5–6
7–812–13
AbsentPresent

Squamation on preopercular flangeStandard length (mm)——Absent?, not observed by enameled
stuff specimen
Absent250

Body depth
Head length
Body width at pectoral fin base
Snout length
Orbit diameter
Interorbital width
Upper jaw length
Caudal peduncle depth
Caudal peduncle length
Predorsal length
Preanal length
Prepelvic length
Dorsal fin baseAnal fin baseCaudal fin length
Pelvic fin spine length
First pelvic fin ray length
Longest pectoral fin ray length
First dorsal fin spine length
Second dorsal fin spine length
Third dorsal fin spine length
First dorsal fin ray length
First anal fin spine length
Second anal fin spine length
Third anal fin spine length
First anal fin ray length

45
32
–
13
9
9
12
10
17
42
70
39
53
20
—
—
—
—
3
5
16 (broken)
—
3
8
8
—

45–46 (46)
31–34 (32)
18–19 (18)
11–17 (13)
10–13 (11)
9–11 (10)11–12 (11)
10–13 (11)
18–20 (19)41–45 (43)65–70 (66)
36–40 (38)
53–58 (56)
21–31 (24)
24–33 (30)
15–17 (16)
19–27 (24)
32–37 (35)
2–5 (4)
5–7 (7)
20–33 (25)
9–11 (11)
4–5 (5)
9–11 (10)
8–11 (10)
8–13 (10)

Scales above and below lateral line
Scales between 5th dorsal fin
spine base and lateral line
Gill rakers
Squamation condition just above eye
Teeth on vomer

1802 in overall body appearance and includes theextended third and fourth dorsal fin spines, butdiffers from the latter in having the fourth dorsal
fin spine being less than 2 times in the fifth dorsal fin spine (vs. over 3 times in E. cardinalis)and scattered cobalt blue spotted markings onsides (vs. several continuous prominent cobalt
blue spotted lines), and usually teeth on vomer

61/2 / 161/2–171/2

Absent126–270 (166)(vs. no teeth) (Tanaka, 1931; Akazaki, 1962;
Carpenter, 2001; this study). Another valid congener, E. ehrenbergii Valenciennes in Cuvier andValenciennes, 1830 is known from the West
African coast and Mediterranean Sea and has aunique character: the third-to-sixth dorsal fin
spines gradually become shorter. Further work is
required for a review of the genus Evynnis.

4–710–11
14–18Large, 2.40–3.35 (2.78)
Shallow, mean 7.8% SL (range
7.6–8.1% SL)
Sharp, 76.9° (70°–83°)a
Rosy to light red with silvery
Reflection, often 7 or 8 wide silver
purplish longitudinal stripes between
each horizontal scale row below
lateral line, visible from lateral view

7–812–13
19–21
Small, 3.09–4.09 (3.66)
Shallow, mean 7.8% SL(6.2–8.8% SL)
Blunt, 80.8° (80°–83°)a
Strong pinkish-red with silvery
reflection with 5 or 6 brightcobalt or blue small-spotted lines(Fig. 2A) between each horizontal
scale row below lateral line,
visible only from oblique dorsal
view, and such lines risingobliquely above lateral line
Uniformly reddish-gold sheen

Pale yellowish with brightyellow distal margin

Uniformly pale

Uniformly pale

Pale yellowish with bright yellow
distal margin

Membrane rays saffron-yellow,
reddish yellow

Strong pinkish-red on most ofpart
Uniformly reddish-yellow
anteriorly and posterodorsally;
posteroventrally hyaline
Mostly strong pinkish-red or
yellow

AbsentAbsentSilvery, sometimes a yellowish
tint ventrally

XIII, 9; III, 8
I, 5; 14–15 (usually 15)
48–50
41/2–5
5–6 / 12–15

Dentex fourmanoiria a
199–233 mm SL (103–226 mm SLa)n316 type specimensa

XII, 10; III, 8
I, 5; 1548–50
41/2–5
5–6 / 12–14, based on Akazaki and Seret (1999).

aColor of caudal fin rays

Color of anal fin rays

Color of pectoral fin rays

Strong yellow blotch on
anterior nostril and snout
regions
Color of dorsal fin rays

Eye color

Snout angle, means (range)
Body color

Orbit diameter in head length
Suborbital depth

Dorsal and anal fin rays
Pelvic and pectoral fin rays
Pored lateral line scales
Scales of transverse series
Scales above and below lateralline
Gill rakers

Dentex abei sp. nov.
128–307 mm SLn40

Mostly saffron yellow or lighter
membrane and rays

Saffron yellow or pale yellow
on most of upper part
Light saffron-yellow
membrane and rays

Saffron-yellow in both
membrane and rays

Generally pinkish-gold or silver
Sheen, sometimes yellowish
tint ventrally
Absent

6–810–13
16–21
Large, 2.37–3.39 (2.78)
Moderate, mean 8.1% SL(range 7.1–10.1)
Sharp, 76.1° (69°–84°)
Light pinkish red with silvery
reflection; wide purplish-silver
longitudinal stripes between eachhorizontal scale row below lateral line

XII, 10; III, 8
I, 5; 15–16
48–50
41/2
5–6 / 13–16

Dentex spariformis
61–283 mm SLn47

Reddish-yellow anteriorly; orangeyellow or yellow posteriorly except
rays and lower margin hyaline
Mostly red to light red;
yellowish anteriorly

Orange-yellow to light red in both
membrane and rays but
longitudinal hyaline stripe
beneath base, especially soft rays
Mostly orange-yellow to reddish

Present

6–9 (usually 7 or 8)10–12(usually 11)16–21 (usually 17–18
Small, 2.66–3.87 (3.32)
Deep, mean 9.5% SL(range 7.3–11.1% SL)
Blunt, 82.9° (82°–88°)a
Red to light red ventrally with
Yellowish-gold sheen, snout vivid
yellow, three large fluorescent yellow
blotches dorsally beneath dorsal finbase plus 1 iris sized-yellow spoton posteriormost soft dorsal fin ray
base (Figs. 1D, 2B), clearly visible
from oblique dorsal view (Fig. 2B)
Uniformly strong reddish gold

XII, 9–10 (usually 10); III, 8
I, 5; 15–16 (usually 15)
47–50
41/2
5 / 13–15

Dentex tumifrons
78–306 mm SL
n54

Table 4. Selected counts, morphometric and color characters of the “Dentex tumifrons complex”: Dentex abei sp. nov., D. fourmanoiri, D. spariformis, andD. tumifrons.

Dentex in Western Pacific45

46

Y. Iwatsuki, M. Akazaki and N. Taniguchi

Remarks. Evynnis japonica Tanaka, 1931
was proposed as a new name for the species
identified and illustrated as Chrysophrys cardinalis of Temminck and Schlegel, 1843 (reproduced as pl. 33; see Fig. 4B). Tanaka (1931) did
not designate a type specimen based on their account on 7 considered to be syntypes by (Boeseman, 1947: 74): RMNH D401 (stuffed, 209 mm
SL), RMNH D402 (stuffed, 253 mm SL), RMNH
D403 (stuffed, 204 mm SL), and RMNH 954a, b,
c, d (4 specimens, 63–143 mm SL). We here des-ignate RMNH 954b (128 mm SL of, Fig. 4C) as
the lectotype of Evynnis japonica Tanaka, 1931,
the other specimens becoming the paralectotypes(Table 3). M. J. P. van Oijen (RMNH) provided
counts and measurements of the lectotype and aparalectotype (RMNH D401) as well as counts of
dorsal and anal fin rays in other paralectotypes
and photographs of all type specimens. As discussed in the treatment of D. hypselosomus, E.
tumifrons is a senior synonym of E. japonica.

Key to Species of the Dentex hypselosomus complex
1a. Dorsal fin with XIII spines and 9 soft rays . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . Dentex fourmanoiri (New Caledonia and Chesterfield Island)
1b. Dorsal fin with XII spines and 10 soft rays. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
2a. Four large fluorescent yellow blotches dorsally below dorsal fin ray base, plus one iris-sized yellow spot on the posteriormost soft dorsal fin ray base (Figs. 1D, 2B), clearly visible from obliquedorsal view (Fig. 2B); snout region prominent vivid yellow . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Dentex hypselosomus (East Asian Shelf)
2b. No large yellow blotches on back; snout without any prominent vivid yellow coloration . . . . . 3
3a. 5 or 6 bright cobalt or blue small spotted lines (Fig. 2A) between each horizontal scale row below
lateral line, such lines rising obliquely above lateral line, easily recognized from oblique dorsal
view (dark brown spots present in preserved specimens); orbit diameter smaller, 3.09–4.09 (mean
3.66 in head length) . . . . . . . . . . . . . . . . . . . Dentex abei sp. nov. (Ryukyu and Ogasawara Islands,
Japan and northeastern Taiwan, plus Luzon Island, Philippines)
3b. No bright cobalt or blue small-spotted lines between each horizontal scale row below lateral line;
orbit diameter larger, 2.37–3.39 in head length (mean 2.78) . . . . . . . . . . . . . . . Dentex spariformis(Australia, except southwestern part, and off Bali and Lombok Island, Indonesia)
Comparative material examined. Evynnis
cardinalis: MUFS 2203, 2204, 2239, 2268, 2306,
5 specimens, 130–146 mm SL, Hong Kong,
China; MUFS 13760, 156 mm SL, Meitsu,
Miyazaki, Japan. E. ehrenbergii: MUFS 22649,
242 mm SL, Canary Islands, Tenerife, Spain.
Acknowledgments
We are most grateful to M. J. P. van Oijen(RMNH), P. Pruvost, G. Duhamel (MNHN), M.
McGrouther (AMS), J. Johnson (QM), H. Larson
and B. Russell (NT), S. Kimura (FRLM), S. Jewett and J. T. Williams (USNM), D. J. Bray and M.

Gomon (NMV), F. Krupp (SMF), M Aizawa(Coastal Branch of Natural History Museum andInstitute, Chiba, Japan) and K. Sakamoto(ZUMT) and for opportunities to examine sparid
type, non-type specimens, and type information,in addition to accommodation and facilities, andalso thank T. Yoshino (URM) for much valuable
historical information, including references and
J. Johnson (QM) for a photograph use of Dentex
spariformis. We thank M. J. P. van Oijen(RMNH) for his translation of Temminck andSchlegel’s article into English and kindly gave 7
syntypes’ photographs of Evynnis japonica andthe lectotype data. Furthermore, we are gratefulDentex in Western Pacificto G. S. Hardy, Ngunguru, New Zealand, for his
critical comments on the initial manuscript andgreat help with English. Lastly, we are very
grateful for many helpful suggestions to improvethe manuscript from two anonymous reviewers
and the editor of K. Matsuura (NSMT). This
study was supported in part by grants awarded to
the first author, by the Ministry of Education,
Science, Sports and Culture, Tokyo, Japan(10640684 and 11640696).
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