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ISSN 0032-9452, Journal of Ichthyology, 2017, Vol. 57, No. 1, pp. 1–9. © Pleiades Publishing, Ltd., 2017.
Samaretta perexilis, a New Genus and New Species of Samarid
Flatfish (Pleuronectiformes: Samaridae) from the South Pacific1
E. P. Voroninaa, * and A. Y. Suzumotob
aZoological Institute, Universitetskaya nab. 1, St. Petersburg, 199034 Russia
bIchthyology, Natural Sciences, Bishop Museum, 1525 Bernice st., Honolulu, HI 96817 United States
*e-mail: voron@zin.ru
Received April 6, 2016
Abstract⎯A new genus of righteye flounder, Samaretta gen. nov., is described from two specimens (one
female and one juvenile) collected in deep waters (470–512 m) from submarine mountains of the southern
eastern Pacific. The type species of the new genus, Samaretta perexilis, is characterized by unremarkable (i.e.,
not elongated) anterior dorsal fin rays, four pectoral fin rays, six parapophyses, reduced lateral line canals, a
very slender body, large head and eyes, and minute scales. An updated key to the samarid genera is presented.
Keywords: Samaretta perexilis, new genus, new species, Samaridae, flatfish, SE Pacific, deep sea
DOI: 10.1134/S003294521701012X
INTRODUCTION
Dwarf righteye flounders of the family Samaridae
have short-based pelvic fins, dorsal fin extending for-
ward on snout below nasal organ of blind side, no pec-
toral fin on blind side, trunk canal usually reduced on
blind side. Among three samarid genera, Samaris
Gray 1831 is characterized by elongate anterior dorsal
fin rays and ocular side pelvic fin rays, pectoral fin
with four rays, unbranched caudal fin rays (Norman,
1934; Venkataramanujam and Ramamoorthi, 1973;
Quéro et al., 1989; Sakamoto, 1984; Hoshino and
Amaoka, 1998). Plagiopsetta Franz 1910 is character-
ized by lacking elongate dorsal and pelvic fin rays, and
possessing a high number of pectoral fin rays (7–10)
(Mihara and Amaoka, 1995). Samariscus Gilbert,
1905 is characterized by 4–6 pectoral fin rays and no
elongate dorsal and pelvic fin rays (Kawai et al., 2011;
Diaz de Astarloa et al., 2013). Samarids live in shallow
to relatively deep waters of the tropical and warm tem-
perate regions of the Indian Ocean and seas of the
western and central Pacific.
Comparative topographic study of the lateral line
canals (LLC) of five flatfish families, including Sama-
ridae, showed stable differences between genera and a
low interspecific variation (Voronina, 2009a), leading
to the conclusion that these peculiarities can be valu-
able diagnostic characters for pleuronectiform genera.
A single specimen (juvenile, 18.3 mm SL) of a
rarely-caught, deep-sea species of Samariscus was
described from Sala y Gomez Submarine Ridge, east-
ern Pacific Ocean (Amaoka et al., 1997). In February
2014, the senior author discovered a non-descript
pleuronectid in the Bishop Museum fish collection
taken by deep dredge in the Tuamotu Archipelago by
Joseph Poupin in October 1990 (Fig. 1). Comparison
of these two specimens revealed their con-specificity.
They share characters mentioned above with
samarids, but body proportions and reduction of LLC
distinguish them from all species of three previously
recognized genera. These specimens are herein
described as a new species and established in a new
genus.
MATERIALS AND METHODS
Two examined specimens are deposited in collec-
tions of the Bernice P. Bishop Museum, Honolulu,
Hawaii, United States (BPBM) and the Zoological
Museum of Moscow University (ZMMU). Counts
and proportional measurements follow Hubbs and
Lagler (1958), except that dorsal and anal fin rays were
counted individually. All measurements were made to
the nearest 0.1 mm with dial calipers. Standard length,
body depth, and head length are abbreviated as SL,
BD, and HL, respectively. Counts of dorsal-, anal-,
and caudal-fin rays, abdominal and caudal vertebrae
and parapophyses of abdominal vertebrae were made
from radiographs. Comparative meristic information
on counts of parapophyses of abdominal vertebrae in
samarids was taken from radiographs of 90 type and
non-type specimens. Comparative information on the
cephalic lateral system of samarid is based on exam-
ination of 55 type and non-type specimens of 21 spe-
cies of three genera, and on previously published data
1The article is published in the original.
2
JOURNAL OF ICHTHYOLOGY Vol. 57 No. 1 2017
VORONINA, SUZUMOTO
(Voronina, 2009a; 2009b). Terminology of the
cephalic lateral system follows that accepted for tele-
osts (Webb, 1989; Mandritsa, 2001) and for pleu-
ronectiforms in particular (Voronina, 2009a). Methy-
lene blue was used in the detection of nostrils, scales
and sensory pores. Institutional abbreviations follow
Sabaj Pérez (Standard Symbolic..., 2014).
FAMILY S AMARIDAE
Samaretta gen. n.
Typ e specie s. Samaretta perexilis sp. n. (Fig. 2)
D i a g n o s i s. The following combination of char-
acters distinguishes Samaretta from other samarid
genera: body slender, BD less than head depth and 3.5
or more in the SL, head large, 3.5 or less in the SL,
eyes large, their diameter less than 3 in HL, anterior
dorsal fin rays very short, their length about 1/2 of eye
diameter; four ocular-side pectoral fin rays; six paired
abdominal vertebrae parapophyses; lateral-line canals
of the head considerably reduced; trunk canals absent
on both sides of body; caudal rays branched except two
uppermost and two lowermost simple, scales minute.
Key to genera of Samaridae
1A Anterior dorsal fin rays greatly elongate and fil-
amentous. Ocular side pelvic fin rays greatly
elongate ………............................. Samaris Gra y, 1831
1B Anterior dorsal and pelvic fin rays not
elongate ................................................................. 2
Fig. 1. Capture locations of holotype (m) and paratype (.) of Samaretta perexilis.
–45°
0°
–15°
–30°
–150 °–165°180 °150 °165°–13 5 °–120°–105°–75°–90°
PACIFIC OCEAN
Fig. 2. Holotype of Samaretta perexilis sp. nov., BPBM 37153, female, 39.0 mm SL, Atoll Mururoa. Scale bar: a, c, d—1 cm, b—
1 mm; nba—anterior nostril of blind side, nbp—posterior nostril of blind side, pt—temporal pore.
(a) (b)
(c) (d)
nba nbp
pt
JOURNAL OF ICHTHYOLOGY Vol. 57 No. 1 2017
Samaretta perexilis, A NEW GENUS AND NEW SPECIES 3
2A Pectoral f in rays 7 or more …Plagiopsetta Franz,
1910
2B Pectoral fin rays 5 or fewer …………………….... 3
3A Depth of body of adults more than head depth
…Samariscus Gilbert, 1905
3B Depth of body of adults less than head depth …..
…................................................ Samaretta gen. nov.
E t y m o l o g y. The new generic name is close to
Samaris Gr ay 1831 a n d Samariscus Gilbert 1905.
R e m a r k s. New genus Samaretta clearly fits in
family Samaridae having eyes on the right side of the
head, no pectoral fin on the blind side, five rays in the
ocular-side pelvic fin. Several external and internal
morphologic features, in particular, remarkable LLC
peculiarities, distinguish Samaretta from three previ-
ously established samarid genera. Stability of the
topography of LLC was demonstrated for a number of
teleost genera (Neelov, 1979; Mandritsa, 2001) as well
as for genera of several pleuronectiform families
(Voronina, 2009a). Lateral line canals were studied in
all samarid species (current data; Voronina, 2009a),
i.e., in five species of Samaris, three species of Plagiop-
setta and 17 species Samariscus. Lateral line canals of
samarids include mandibular canal (two sensory
pores), preopercular canal (5–6 pores), temporal
canal, supratemporal canal (2–3 pores) on both sides
of head and supraorbital canal of the ocular side (two
pores) (Fig. 3). Infraorbital canal is absent on both
sides of head. Coronal commissure connects canals of
both sides. Trunk canal is usually present on the eyed
and absent on the blind side, in some species it is pres-
ent (Mihara et al., 2004) or absent on both sides
(Voronina, 2009b). Intergeneric differences concern
supraorbital canal of blind side – this canal with one-
two pores is present in Samariscus and absent in
Samaris and Plagiopsetta. Reduction of LLC distin-
guishes Samaretta from other samarids: temporal
canal of blind side reduced to one pore; temporal
canal of the eyed side, supratemporal and trunk canals
absent on both sides; there are only four pores in
shorter preopercular canal on the blind side. Absence
of trunk canal on both sides and pores above nostrils
on the blind side was mentioned in paratype descrip-
tion (Amaoka et al., 1997). Reduction of temporal
canal through frontale, pteroticum, supratemporale
and posttemporale is rare in teleosts and, according to
our preliminary data, unique for pleuronectoids.
Interspecific differences of caudal vertebrae num-
ber are traditionally used in diagnoses of samarid spe-
cies. Number of vertebrae of Samaretta (10+30–32) is
within variation for other samarids: 9–10+27–37 in
Samariscus (Kawai et al., 2011), 9–11+29–35 in
Samaris (Diaz de Astarloa et al., 2013) and 10–11+27–
34 in Plagiopsetta (Mihara and Amaoka, 2004). Other
vertebral characters, e.g., presence of parapophyses
(=transverse apophyses) were recorded for some pleu-
ronectiforms including samarids (Clothier, 1950;
Amaoka, 1969; Hensley and Ahlstrom, 1984; Saka-
moto, 1984). However, these findings were not ana-
lyzed and used in flatfish taxonomy. Our study of
radiographs of samarids finds significance in different
numbers of parapophyses, occurring as one pair on the
anterior end of several abdominal vertebrae. There are
six pairs of parapophyses in both specimens of Sama-
retta and in five specimens of three Plagiopsetta spe-
cies; six or seven pairs of parapophyses in 39 speci-
mens of five Samaris species (Table 1, Fig. 4). There
are only five pairs of parapophyses in all 44 specimens
of 19 species of Samariscus. Five parapophyses is a
synapomorphy of Samariscus species vs six or more
parapophyses in Samaretta, Samaris, Plagiopsetta.
Therefore this character distinguishes Samaretta from
Samariscus. Moreover, characters of external mor-
phology—an extremely slim body, very large head, and
relatively large eyes also distinguish S. perexilis from all
presently known valid species of the genus Samariscus
(Table 2). General differences of BD, head and eye
size used in identification keys for Samariscus species
with comparison to Samaretta are summarized in
Table 3. Moreover, BD is noticeably less than head
depth in Samaretta, whereas it exceeds head depth in
all Samariscus species (Fig. 5). Eye diameter of Sama-
retta compared with head length is within intraspecific
variation demonstrated for Samariscus species (Table 2).
However eye size compared with body length is con-
siderably larger (8–10 vs 13–20 in SL in Samariscus
species). Scales of Samaretta are minute, not overlap-
ping, half the size on Samariscus and Samaris of com-
parable body length (Fig. 6).
Non-elongate anterior rays of dorsal fin in an adult
specimen and branched midcaudal rays unambigu-
ously distinguishes Samaretta from Samaris; presence
of only four rays in pectoral fin—from Plagiopsetta.
Therefore, a combination of unique topography of
the LLC, six pairs of abdominal vertebrae parapophy-
ses, four rays in pectoral fin, anterior rays of dorsal fin
not elongate, branched midcaudal rays, distinctive
proportions of the body, head, and eyes, and very
small size of scales provide the basis for the assignment
of new genus Samaretta. Futher support is provided by
scleral ossicle counts. Consistency of scleral ossicle
numbers in teleost families, and, in particular, varia-
tion from zero to two in Pleuronectiformes, has been
demonstrated (Franz-Odendaal, 2008; Mok and Liu,
2012; Voronina and Chanet, 2014). Absence of scleral
ossicles in both specimens of Samaretta as opposed to
Table 1. Distribution of paraphophyses number of samarid
genera
Samarid genera Paraphophyses number
55 66 7
Samariscus (19 species) 44
Samaretta (one species) 2
Samaris (five species) 14 25
Plagiopsetta (three species) 5
4
JOURNAL OF ICHTHYOLOGY Vol. 57 No. 1 2017
VORONINA, SUZUMOTO
two scleral ossicles in other specimens in the current
study distinguishes Samaretta from three other
samarid genera.
The only known species of Samaretta is from sub-
marine mountains in the southern eastern Pacific.
Samaretta perexilis sp. nov. (Fig. 2)
Samariscus sp. Amaoka et al., 1997
H o l o t y p e: BPBM 37153, 39.0 mm SL, gravid
female, 21o51′S 139o1′W, Tuamotu Archipelago Atoll
Fig. 3. Lateral line canals of the head of samarid genera: (a) Samariscus, (b) Samaris and Plagiopsetta, (c) Samaretta. CMb—man-
dibular canal blind side, CM—coronal commissure, CMo—mandibular canal ocular side, CPb—preopercular canal blind side,
CPo—preopercular canal ocular side, CSOb—supraorbital canal blind side ocular side, CSOo—supraorbital canal ocular side,
CSTb—supratemporal canal blind side, CSTo—supratemporal canal ocular side, CTb —temporal canal blind side, CTo—temporal
canal ocular side.
CMb
CPb
CSOb
CTb CMC
CSOo
CSTb CSTo
CTo
CMo
CPo
(a)
(b)
(c)
JOURNAL OF ICHTHYOLOGY Vol. 57 No. 1 2017
Samaretta perexilis, A NEW GENUS AND NEW SPECIES 5
Mu ruro a, d epth 512 m, d redge, colle cted by J. Po upin,
17 October 1990.
P a r a t y p e: ZMMU 39939, 18.3 mm SL, 25o S
98o W, depth 470 m, unnamed seamount of the Sala y
Gomez Submarine Ridge, R/V Professor Shtokman,
cruise 18, station 1995, collected by N.V. Parin and
colleagues, May 4, 1987.
Diagnosis. As for genus.
D e s c r i p t i o n. Counts and proportional mea-
surements (% SL) of the holotype are listed first, fol-
lowed by those of the paratype placed within paren-
theses.
Dorsal fin rays 69 (64); anal f in rays 54 (49); ocu-
lar-side pectoral fin rays 4 (4), pectoral fin absent on
blind side; pelvic fin rays on ocular side 5 (5) and on
blind-side 4 (5); caudal fin rays 16 (16); abdominal
vertebrae 10 (10), caudal vertebrae 32 (30); total verte-
brae 42 (40).
Head length 28.5 (32.0) % SL; body depth 24.6
(29.8); snout length 6.4 (6.7); upper eye diameter 10.0
(12.4); lower eye diameter 10.0 (12.4); interorbital
width 1.3 (1.7); upper jaw length 11.0 (8.9) on ocular
side, 10.2 (9.6) on blind side; lower jaw length 14.9
(14.6) on ocular side, 14.9 (15.2) on blind side; depth
of caudal peduncle 12.0 (11.2); pectoral fin length 21.8
(17.4) on ocular side; pelvic fin length 8.7 (7.9) on
ocular side, 5.9 (5.1) on blind side; length of caudal
peduncle 3.9 (3.9).
Body slender. Eyes dextral, separated by narrow,
scaleless bony ridge. Mouth oblique, moderate size,
almost symmetrical, maxilla extending below anterior
margin of lower eye. Snout short, smaller than either
eye diameter. Nostrils on ocular side open at tip of
Fig. 4. Radiographs of anterior part of body showing the number of transverse apophyses of abdominal vertebrae (arrows) in four
samarid species. (a) Samaretta perexilis (BPBM 37153, holotype 39.0 mm SL), (b) Samaris cristatus (ZIN 48920, 119.4 mm SL),
(c) Plagiopsetta stigmosa (ZIN 55366, former MNHN 2000–5483, paratype 143.8 mm SL), (d) Samariscus luzonensis (USNM
93098, holotype 57.1 mm SL); so—scleral ossicles.
(a)
(b)
(c)
(d)
so
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JOURNAL OF ICHTHYOLOGY Vol. 57 No. 1 2017
VORONINA, SUZUMOTO
tubes: anterior tube long; posterior tube very short.
Nostrils on blind side located ventrally to dorsal fin
origin, very obscure, without tube. Teeth small, coni-
cal, densely arranged in bands, equally developed on
jaws of both sides. Vomer and palatines toothless. Gill
rakers on first arch rudimentary on both sides. Gill
membranes united to each other, but not to body.
Upper eye close to dorsal margin of head, on the same
vertical line or slightly in advance of lower eye. Eye
diameter 1/3 of the head length. Snout somewhat
rounded; shorter than eye diameter. Scales minute,
not overlapping, cycloid or weakly ctenoid on ocular
and cycloid on blind side of body of holotype, absent
on snout and opercular regions. Trunk canal absent on
both sides of body; there are only nine sensory pores
on the ocular side of head and seven pores on the blind
side (Fig. 3c). Dorsal-f in origin anterior (holotype) or
slightly posterior (paratype) to anterior margin of
upper eye. Anal-fin origin just posterior to anus. All
dorsal and anal rays simple, none elongate. First and
second pectoral fin rays on ocular side a little longer
than other two, all rays simple; no blind side pectoral
fin. Pelvic fins small, subsymmetrical; first ray of
blind-side fin opposite space between first and second
ray of ocular-side fin; all rays simple. Caudal peduncle
very short, rather deep. Caudal fin rounded, inner 12
caudal rays branched, two dorsalmost rays and two
ventralmost rays simple. Genital papilla on ocular side
posterior to pelvic fin. Vent opening just posterior to
pelvic fin on midventral line. Hypomerals only in
abdominal region. Epimerals and epicentra not visible
in radiograph. Hypurals 2, 3 and 4 fused to first preu-
ral centrum; hypurals 1 and 5 autogenous.
Color in alcohol. According to published
data for freshly-caught paratype there are four and five
dark blotches on the ocular side along the dorsal and
ventral margins of the body respectively; entire pecto-
ral fin darkly pigmented (Amaoka et al., 1997). Pres-
ent pigmentation of both specimens completely lack-
ing on both sides.
S e x u a l d i m o r p h i s m. The large eggs
(approximate diameter 0.5–1.0 mm) clearly seen
through transparent wall of body of holotype leave no
doubt that this specimen is the adult female (Figs. 2a
and 2b). The paratype is very small (17.8 mm SL) and
the sex can not be determined. No known males are
available.
D i s t r i b u t i o n. Known only from submarine
mountains of the southern eastern Pacific, at 470–512
m depth.
Etymology. The name perexilis, from Latin,
references the very lean body.
Remarks. The paratype of S. perexilis was earlier
described as a juvenile Samariscus sp. (Amaoka et al.,
1997), plausibly representing an undescribed species
requiring adult specimens from the same general
vicinity for clarification. Some meristic features of
S. perexilis lie within intraspecific variation of several
Samariscus species. However, the chance discovery of
an adult female pleuronectid of uncertain genus from
the same general area sharing osteological characters
along with LLC, body, and scale characteristics
unique to these specimens argues firmly for the con-
clusions set forth presently.
Samarid species are distributed in tropical waters of
the Indo-west Pacific from South Africa and Mada-
gascar to the Hawaiian Islands and New Caledonia
(Quéro et al., 1989; Kawai et al., 2011). The Eastern
Pacific barrier, which has prevented 90% of all Indo-
Pacific fishes from reaching the west coasts of South
and Central America has also prevented samarids from
establishing themselves there. Samaretta perexilis
occurrences, very distant from other samarid ranges,
extend the longitudinal limit of this family eastward by
about 60 degrees.
Comparative materials. Plagiopsetta glossa, ZIN
36127, 1 specimen, 116.9 mm SL, East China Sea,
28o38′N, 126o30′S, x-ray, USNM 93090, holotype of
Samariscus fasciatus, junior synonym of P. glossa,
South China Sea, Hong Kong, x-ray only; Plagiopsetta
gracilis, MNHN 2000–0706, holotype, 40.5 mm SL,
New Caledonia, x-ray; Plagiopsetta stigmosa, MNHN
2014–2550, 2014–2565, 2014–2392, 5 specimens,
97.2–151.3 mm SL, New Caledonia, ZIN 55366 (former
MNHN 2000–5483), 2 paratypes, 98.4 and 143.8 mm
SL, New Caledonia, x-ray; Samaris chesterfieldensis,
MNHN 2000–0737, 2000–0738, 3 paratypes, 48.5–
50.8 mm SL, Chesterfield Islands, ZIN 55367 (former
Table 2. Measurements (mm) of the holotype and paratype
Samaretta perexilis
Measurements, mm Holotype
Paratype
Amaoka et al.,
1997 /present
study
Standard length 39.0 18.3/17.8
Body depth 9.6 5.7/5.3
Head length 11.1 5.7/5.7
Snout length 2.5 1.2/1.2
Upper eye diameter 3.9 2.2/2.2
Lower eye diameter 3.9 2.2/2.2
Interorbital width 0.5 na/0.3
Upper jaw length on ocular side 4.3 1.8/1.6
Upper jaw length on blind side 4.0 1.9/1.7
Lower jaw length on ocular side 5.8 2.8/2.6
Lower jaw length on blind side 5.8 2.7/2.7
Pectoral fin length on ocular side 8.5 3.4/3.1
Pelvic fin length on ocular side 3.4 0.7/1.4
Pelvic fin length on blind side 2.3 0.4/0.9
Depth of caudal peduncle 4.7 2.2/2.0
Length of caudal peduncle 1.5 0.5/0.4
JOURNAL OF ICHTHYOLOGY Vol. 57 No. 1 2017
Samaretta perexilis, A NEW GENUS AND NEW SPECIES 7
MNHN 2000–0737), 1 paratype, 51.7 mm SL, Ches-
terfield Islands, x-ray; Samaris costae, ZIN 48926, 1
specimen, 35.8 mm SL, Indian ocean, Saya de Malha
Bank, x-ray, ZIN 48927, 1 specimen, 104.4 mm SL,
Indian ocean, Saya de Malha Bank, x-ray, ZIN 48928,
2 specimen, 92.8 and 95.4 mm SL, x-ray, Indian
ocean, Saya de Malha Bank, ZIN 49501, 2 specimens,
27.4 and 47.0 mm SL, x-ray, Indian Ocean, 3o51′ S,
56o08′ E; Samaris cristatus, ZIN 48919, 1 specimen,
131.6 mm SL, Indian ocean, 7°56′ N, 77°03′ E, x-ray,
ZIN 48920, 3 specimens, 119.4–126.8 mm SL, Indian
ocean, 8°48′ N, 76°17 ′ E, x-ray, ZIN 54013, 1 speci-
men, 32.6 mm SL, Gulf of Tonkin, x-ray, ZIN 55101,
1 specimen, 91.1 mm SL, Gulf of Tonkin, x-ray, ZIN
55900, 1 specimen, 110.5 mm SL, Gulf of Tonkin, x-
ray, ZIN 55551–55562, 56025, 24 specimens, 31.1–
126.7 mm SL, Nha Trang Bay, Vietnam, x-ray;
Samaris macrolepis, BMNH 1888.12.1.32, holotype,
34.3 mm SL, Gulf of Martaban, x-ray, HUMZ-L
00896, 9.7 mm SL, x-ray only, 19°59′S, 117°3′ E;
Samaris spinea, MNHN 2000–0724, holotype, 48.7
mm SL, New Caledonia, MNHN 2000–0723, para-
type, 46.9 mm SL, New Caledonia, ZIN 55368 (for-
mer MNHN 2000–0716, 2000–0717), 2 paratypes,
31.8 and 33.9 mm SL, Chesterfield Islands, x-ray, ZIN
54013, 1 specimen, 32.6 mm SL, Pacific Ocean,
29°29′ S, 167°52′ E, x-ray, ZIN 55281, 1 specimen,
35.1 mm SL, Coral Sea 25°14′ S, 159°48′ E, BPBM
39606, 29°29′S, 167°52′E,Chesterfield Islands;
Samariscus corallinus, USNM 51596, holotype,
Hawaiian Archipelago, Oahu, x-ray only, USNM
51676, 2 paratypes, Hawaiian Islands, Oahu, x-ray
only ZIN 51735, 4 specimens, 84.9–148.0 mm SL,
Pacific Ocean, 25°31′ N, 169°35′ W, x-ray, BPBM
Fig. 5. Proportions of the head, body and eyes of samarids:
(a) Samaretta perexilis, (b) Samariscus maculatus, (c)
Samariscus sunieri, (d) Samariscus latus.
(a)
(b)
(c)
(d)
Table 3. Characteristics of body, head and eyes of Samaretta perexilis and Samariscus species
Generalized information for Samariscus species from our data; Norman, 1934; Woods, 1966; Ochiai and Amaoka, 1962; Shen, 1982;
Quéro et al., 1989; Kawai et al., 2008; Kawai et al., 2011.
Object Body
(depth in SL)
Head
(length in SL)
Eye
(lower eye diameter
in head length/in SL)
Samaretta perexilis slender (3.5 to 4) very large (3 to 3.5) large (2.6 to 2.8/8 to 10)
Samariscus species elongate (2.5 to 3.4) large (3.5 to 3.7) moderate (2.6 to 3.5/13–17)
elliptical (2.1 to 2.5) small (4.0 to 5.6) small (4/20)
Fig. 6. An ocular-side scale of samarids: (a) Samaretta per-
exilis (BPBM 37153, 39.0 mm SL), (b) Samariscus multira-
diatus (MNHN 2005–0015, 38.5 mm SL), (c) Samaris
spinea (ZIN 55368, former MNHN 2000–0716, 34.1 mm
SL). Scale bar: 1 mm.
(a) (b) (c)
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JOURNAL OF ICHTHYOLOGY Vol. 57 No. 1 2017
VORONINA, SUZUMOTO
15507, 1 specimen, 78.4 mm SL, Hawaiian Islands,
Oahu; Samariscus desoutterae, MNHN 1984–0368,
holotype, Madagascar, x-ray only, MNHN 1984–
0369 paratype, 79.5 mm SL, Madagascar, x-ray;
Samariscus filipectoralis, ZIN 55548, 1 specimen,
90.9 mm SL, South China Sea, off Vietnam, x-ray;
Samariscus hexaradiatus MNHN 2002–3655, holo-
type, 100.3 mm SL, Solomon Islands, MNHN 2002–
3671, paratype, 80.9 mm SL, Solomon Islands, x-rays;
Samariscus huismani, MNHN 2009–1562, 4 speci-
mens, 65.2–83.4 mm SL, Coral Sea, NE Tutuba
Island, MNHN-IC-2012-0805, 6 sp., Coral Sea,
Tutuba Island, Vanuatu Archipelago, x-ray only;
Samariscus inornatus, BMNH 1927.1.6.67, syntype,
96.5 mm SL, Gulf of Aden, x-ray; Samariscus japoni-
cus, MNHN-IC-1992-0936, Tosa bay, Kochi prefec-
ture, Japan, x-ray only; Samariscus latus, ZIN 48921,
1 specimen, 78.7 mm SL, Saya de Malha Bank, ZIN
48922, 2 specimens, 77.5 and 78.6 mm SL, Saya de
Malha Bank, x-ray, ZIN 48923, 3 specimens, 74.9–
82.9 mm SL, Saya de Malha Bank, x-ray, ZIN 54700,
1 specimen, 83.5 mm SL, x-ray, Saya de Malha Bank;
Samariscus leopardus, ZMMGU P-15039, holotype,
122.0 mm SL, Saya de Malha Bank, x-ray; Samariscus
longimanus, BMNH 1927.1.6.68, paratype, 95.4 mm
SL, West of Ceylon, USNM 137384–137388, 8 speci-
mens, Philippines, x-rays only; Samariscus luzonensis,
USNM 93089, holotype, Philippines, West coast of
Luzon, Manila Bay to Lingayen Gulf, x-ray only, ZIN
555549, 1 specimen, 32.3 mm SL, Nha Trang Bay,
Vietnam, x-ray, ZIN 55550, 1 specimen, 50.7 mm SL,
Nha Trang Bay, Vietnam, x-ray, ZIN 55792, 1 speci-
men, 52.6 mm SL, Nha Trang Bay, Vietnam, x-ray,
ZIN 56026, 1 specimen, 48.8 mm SL, x-ray, Nha
Trang Bay, Vietnam, ZIN 55901, 1 specimen,
39.0 mm SL, Gulf of Tonkin, x-ray, ZIN 55902,
1 specimen, 46.0 mm SL, Gulf of Tonkin, x-ray, ZIN
55903, 1 specimen, 41.2 mm SL, Gulf of Tonkin, x-
ray; Samariscus macrognathus, USNM 93088, holo-
type, Philippines, West coast of Luzon, Manila Bay to
Lingayen Gulf, x-ray only, MNHN 2006–0195,
2 specimens, 64.2 and 85.3 mm SL, Solomon Islands,
8°36′ S, 157°23′ E; Samariscus maculatus, BMNH
1879.5.14.84, holotype, 81.6 mm SL, Kai Is., Arafura
Sea, x-ray; Samariscus multiradiatus MNHN-IC-
2006-1758, holotype, New Caledonia, x-ray only,
MNHN 2005–0015, 5 specimens, 38.3–45.6 mm SL,
New Caledonia; Samariscus neocaledonia MNHN-
IC-2011-0231, holotype, 48.3 mm SL, New Caledo-
nia, x-ray; Samariscus nielseni MNHN-IC-1984-
0367, holotype, Madagascar, x-ray only, MNHN
2006-0196, 1 specimen, 77.8 mm SL, Solomon
Islands, 8°36′ S, 157°23′ E; Samariscus sunieri,
BMNH 1933.2.18.17 syntype, 128.0 mm SL, St. Niko-
las Bay, Bali, x-ray; Samariscus triocellatus, USNM
141767, holotype, Marshall Islands, x-rays only,
USNM 141768, 3 paratypes, Marshall Islands, x-rays
only, BPBM 28152, 3 specimens, 64.7–77.3 mm SL,
ZIN 55538 (former BPBM 28152), 1 specimen,
54.5 mm SL, Hawaiian Islands, Oahu, x-ray;
Samariscus xenicus, MNHN-IC-1992-0937, Tosa
Bay, Kochi prefecture, Japan, x-ray only.
ACKNOWLEDGMENTS
We would like to thank E. Vasil’eva (ZMMU), for
the loan of the Sala y Gomez paratype and V. Sideleva
and A. Balushkin (ZIN), for discussion and critique of
the manuscript. The authors also extend sincere grati-
tude to these colleagues for their kind help and provid-
ing radiographs: P. Pruvost, R. Causse, Z. Gabsi and
C. Ferrara (MNHN), J. Maclaine and N. Martin
(BMNH), H. Imamura and S. Ohashi (HUMZ), J.-
M. Diaz de Astarloa (Universidad Nacional de Mar
del Plata, Argentina) and USNM Ichthyological
Department staff, especially J. Williams and S. Rar-
edon.
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