Bull. Natl. Mus. Nat. Sci., Ser. A, Suppl. 4, pp. 49–66, March 29, 2010

Myopsaron nelsoni, a New Genus and Species of Sandburrowers(Perciformes: Trichonotidae: Creediinae) fromthe Ogasawara Islands, Japan
Koichi Shibukawa
Nagao Natural Environment Foundation, 3–10–10 Shitaya, Taito-ku, Tokyo 110–0004, Japan
E-mail: kshibukawa@nagaofoundation.or.jp

Abstract A new genus and species of the trichonotid subfamily Creediinae, Myopsaron nelsoni,
is described based on 43 specimens, including 20 type specimens (22.1–36.3 mm in standard
length), collected by sledge nets at the depths of 55–99 m off southern Chichi-jima Island,Ogasawara Islands, Japan. It is readily distinguished from the other genera of the subfamily byhaving some unique, apparently autapomorphic, features, e.g., the globular fleshy extension at the
tip of upper jaw, discontinuous lateral line on body, higher number of precaudal vertebrae thancaudal vertebrae (24–2618–2143–45 vertebrae), and edentate premaxilla. Myopsaron is highly
specialized, but, as well as Creedia, it appears to be nested within the basal creediines since it lacks
several specializations found in all other creediines exclusive of the most primitive member,
Schizochirus.
Key words : Myopsaron nelsoni, New genus and species, Creediinae, Ogasawara Islands.

Fishes of the Indo-Pacific trichonotid subfamily Creediinae (sensu Smith and Johnson, 2007)
are small, slender sandy-bottom dwellers, comprising about 16 species belonging to the following seven genera (Nelson and Randall, 1985;
Yoshino et al., 1999; Nelson, 2006): Apodocreedia, Chalixodytes, Creedia, Crystallodytes,
Limnichthys, Schizochirus, and Tewara. Althoughsimilar to their allies (i.e., trichonotines andhemerocoetines), the creediines are readily distinguished from the other trichonotids by several
characters, e.g., upper jaw with fleshy extension,slightly protruding eye with infolding of cornea
at orbit margin, and lateral line descending
abruptly or gradually to ventral portion of caudal-fin base (Nelson, 1985). The creediines areprimarily found on the flat, fine sandy bottoms
of shallow coastal waters, but some (e.g.,
Limnichthys rendahli and Creedia haswelli) havebeen captured from more than 100 m depth (Nelson, 1978, 1983).
During a biological survey of the Ogasawara
Islands and the adjacent waters carried out by
TR/V Seiyo-maru of Tokyo University of Fish-eries (now Tokyo University of Marine Scienceand Technology) in June 1995, 43 specimens ofan unusual creediine fish (Figs. 1 and 2) were
captured by sledge nets at depths of 55–99 m off
the southern coast of Chichi-jima Island. Subsequently, a graduate-course student of the university on the cruise, T. Kanazawa (now at Oita Prefecture, Japan), kindly provided me an opportunity to examine these specimens. My research
revealed that the specimens represented a new
species that could not be placed in any existing
genera. The creediine fish, herein described as
new genus and species, is readily distinguishedfrom all known members of the subfamily by
several unique characters, i.e., globular-shaped
fleshy extension at the tip of upper jaw, discontinuous lateral line on body, higher number of
precaudal vertebrae than caudal vertebrae (viz.
24–2618–2143–45 vertebrae), and edentate
premaxilla.
This species has been noticed briefly by Shimada (2002: 1588). He noted “Some specimens . . . have been collected from Ogasawara
and Ryukyu Islands,” but mention of the Ryukyu

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Koichi Shibukawa

Islands was in error, and the species is hitherto
known only by the Ogasawara specimens noted
above.
Materials and Methods
Institutional abbreviations follow Leviton etal. (1985). All fish lengths given are standard
lengths (SL). Measurements were made withcalipers under dissecting microscopes to thenearest 0.01 mm. The methods for measurements
followed those of Hubbs and Lagler (1958), withexceptions given below (the snout tip refers to
the mid-anteriormost point of the fleshy extension at the anterior tip of the upper jaw): body
depth was measured at the dorsal-fin origin; pectoral- and pelvic-fin lengths were represented by
the length of the longest ray of the respective fin;preanal and prepelvic length were measured fromthe snout tip to the base of the first segmented
anal-fin ray and pelvic-fin spine, respectively;
preanus length was measured from the snout tip
to the anus; caudal-peduncle depth was measured
at the anterior base of the anteriormost procurrent ray of the caudal fin. All fin ray elements
were counted, including the uppermost rudimentary ray of the pectoral fin and the ultimate simple ray of the dorsal and anal fins. Count of longitudinal scales is the number of oblique scalerows (anterodorsal to posteroventral) and is takenfrom the anteriormost lateral-line scale posteriorly to the mid-base of the caudal fin (posterior
margin of hypural plate). Transverse scales werecounted in two ways: upper transverse scales are
the number of scale rows between the lateral line
and the origin of the dorsal fin; lower transverse
scales are the number of scale rows between thelateral line and the origin of the anal fin. Predorsal scales included all scales along the pre-dorsal
midline. Gill rakers included the rakers on theouter surface of the first arch including all rudiments; they are given in the text in the form of
the rakers on the upper arch (epibranchial 1)
lower arch (ceratobranchial 1hypobranchial1). Osteological features were observed fromradiographs and specimens cleared and counterstained for bones and cartilages, following themethods of Dingerkus and Uhler (1977) or Potthoff (1984); precaudal vertebrae are the anteriorvertebrae lacking a distinct haemal spine, whereas the others (including urostylar complex) werecounted as caudal vertebrae. Osteological terminology followed those of Suda (1991, 1996), except as follow: intermuscular bones were identified as epineurals rather than epipleurals, following Johnson and Patterson (1993) and Patterson
and Johnson (1995); anteriormost element of ethmoid region of skull was identified as vomerrather than prevomer; nomenclature of caudal
skeleton elements followed that of Fujita (1989,1990). Teeth on the lower jaw of the new species
were quite minute and difficult to discern, andwere confirmed in the cleared and stained specimens. Cephalic sensory canals were observed on
the specimens stained with cyanine blue (seeSaruwatari et al., 1997). Sex was determined primarily externally (see “Sexual dimorphism” andFig. 4), and, in some specimens, verified by internal examination. Observations and illustrationsof osteological features and cephalic sensory systems were made using a dissecting microscopewith a camera lucida. All photographs and line
drawings were made by the author.
Myopsaron gen. nov.
[New Japanese name: Nezu-sunaginpo zoku]

Type species. Myopsaron nelsoni Shibukawa, new species.
Diagnosis. The new genus Myopsaron is
unique amongst the creediines in having: globular fleshy extension at tip of upper jaw; lateralline on body divided into 2 parts, with 3 (rarely 2or 4) grooved scales in anterior series and 33–35
pored scales with distinct posterior lobe in posterior series; higher counts of precaudal vertebrae
than caudal vertebrae (vertebral counts 24–26
18–2143–45); edentate premaxilla. In addition,
it is characterized by the following features:
14–16 dorsal-fin rays; 16–18 anal-fin rays; dorsal-plus anal-fin rays 30–34; both dorsal and anal
fins restricted to posterior half of body; anal-fin

New Creediine Fish from Ogasawara Islands

origin below first to third dorsal-fin rays; I, 3–4(almost always I, 3) pelvic-fin rays; fourth pectoral- and first pelvic-fin rays greatly elongatedand filamentous in male; all dorsal-, anal-, pectoral- and pelvic-fin segmented rays unbranched;
9 branched caudal-fin rays; 1 (rarely two) anal-fin
pterygiophores anterior to first haemal spine; noteeth on vomer and palatine, 1 to 3 minute conical teeth on middle of dentary; 2 epurals; bothanterior and posterior nares pore-like; suborbitalskin fold discontinuous at midway; infraorbital
canal shortly interrupted below eye; mandibular
canal not joined with preopercular canal.
Comparison. Myopsaron is readily distinguished from the other creediines by the diagnostic characters listed above, particularly the globular fleshy extension at the tip of the upper jaw(vs. beak-like fleshy extension along most oflength of upper jaw in the other creediids) anddiscontinuous lateral line on body (vs. continuous). Myopsaron is similar to Creedia in several
meristic counts (e.g., number of fin rays, longitudinal scales, vertebrae and epurals) and coloration, but Creedia possesses vomerine teeth aswell as a continuous lateral line and beak-likefleshy extension on the upper jaw (Fig. 5B).
Etymology. The new generic name Myopsaron
is derived from the combination of the Greek
words mys, meaning “mouse,” and opsaron, aterm for a little fish, in reference to its uniqueglobular-shaped fleshy extension at tip of upper
jaw, resembling the nose of mouse. The gender isneuter.

51

Myopsaron nelsoni sp. nov.
[New Japanese name: Nezu-sunaginpo](Figs. 1–4, 5C, 6–9, 10A and 11–20; Tables 1 and 2)
Holotype. NSMT-P 94886, 35.6 mm SL, male, offsouthern part of Chichi-jima Island, Ogasawara Islands,
Japan (27°1.80N, 142°12.22E–27°1.87N, 142°12.38
E), 51–68 m depth, sledge net, TR/V Seiyo-maru, 19 June
1995.
Paratypes. Total 19 specimens, 22.1–36.3 mm SL:
AMS I.44840-001, 3 specimens (2 males and 1 female),
34.3–36.3 mm SL, collected with holotype; NSMT-P
94887, 8 specimens (7 males and 1 female), 33.8–36.3mm SL, off southern Chichi-jima Island, Ogasawara
Islands, Japan (27°1.74N, 142°12.48E–27°1.78N,
142°12.62E), 95–99 m depth, sledge net, TR/V Seiyomaru, 19 June 1995; NSMT-P 94888, 2 specimen (1 maleand 1 female: cleared, stained and dissected), 32.5–34.6mm SL, same collecting data with holotype; NSMT-P
94895, 1 specimen (male, stained with alizarin red), 31.8mm SL, collected with NSMT-P 94887; OMNH-P
35374–35376, 3 specimens (2 males and 1 female), 22.1–
26.8 mm SL, collected with holotype; YCM-P 44207, 2
specimens (males), 33.1–33.4 mm SL, collected withholotype.
Non-type materials. Total 23 specimens, 16.6–22.7mm SL: NSMT-PL 722, 4 specimens, 16.8–22.7 mm SL,
collected with NSMT-P 94887; NSMT-PL 723, 17 specimens, 16.6–19.4 mm SL, collected with holotype; NSMTPL 724, 2 specimens (cleared and stained), 16.6–18.2 mm
SL, collected with holotype.

Diagnosis. See generic diagnosis.
Description. In the following description,
the counts of the holotype are asterisked, and thefrequency of each count is given in parentheses
following the relevant count. Dorsal-fin rays 14(3), 15* (7) or 16 (7); anal-fin rays 16 (4), 17*(12) or 18 (1); dorsal- plus anal-fin rays 30 (1),31 (3), 32* (8), 33 (4) or 34 (1); anal- minus dorsal-fin rays 0 (2), 1 (5), 2* (8) or 3 (2); pectoral-

Fig. 1. Myopsaron nelsoni sp. nov., 1 of AMS I.44840-001, 34.3 mm SL, male, off southern Chichi-jima Island,Ogasawara Islands, Japan.

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Koichi Shibukawa

Fig. 2. Dorsal (A) and lateral (B and C) views of Myopsaron nelsoni sp. nov., 2 of AMS I.44840-001 in 70%
ethanol-preserved condition. A–B) male, 34.3 mm SL; C) female, 36.3 mm SL.

fin rays 11 (4), 12* (23) or 13 (5); pelvic-fin rays
I, 3 (30) or I, 4 (1); branched caudal-fin rays
54* (17); dorsal unbranched caudal-fin rays 2(5) or 3* (11), all unsegmented except for posterior most 1 (16) segmented ray; ventral unbranched caudal-fin rays 4* (15) or 5(1), all unsegmented except for posteriormost 1* (16) segmented ray; lateral line discontinuous, with 2 (1),3* (29) or 4 (1) grooved scales in anterior series
located above pectoral-fin base, and 33 (12), 34*(19) or 35* (2) pored scales with distinct posterior lobe in posterior series running along ventral
profile of body; longitudinal scales 42 (3), 43(17), 44* (8) or 45* (1); predorsal scales 24 (3),
25 (8), 26* (4), 27 (1) or 28 (1); upper transverse
scales 3* (32); lower transverse scales 1* (30) or
2 (2); gill rakers on outer surface of first arch09 (4), 010 (5), 011* (15) or 012 (6);pseudobranchial filaments 3* (27) or 4 (1).
Proportional measurements are given in Table
1. Body slender, cylindrical, somewhat compressed posteriorly. Snout pointed, gently slopingin dorsal profile. Upper jaw subequal to, and notprojecting beyond, lower jaw, with a globular

fleshy extension at anterior tip (Figs. 1, 3 and 7).
No teeth on premaxilla, vomer and palatine; dentary with 1–3 minute conical teeth. Maxilla extends posteriorly to beyond anterior margin ofeye. Lower jaw bordered by uniserial row ofshort cirri, 10–14 (13 in holotype) per side.
Tongue slender with pointed anterior tip, freefrom floor of mouth. Eye slightly protruding, as
does the lens, with infolding of cornea aroundlens margin; eye without dorsal iris flap. Suborbital skin fold slightly covers lower margin ofeye, discontinuous at midway (Figs. 3 and 7).
Both anterior and posterior nares pore-like. All
dorsal-, anal-, pectoral- and pelvic-fin rays segmented and unbranched. Anal fin originates at avertical through base of first to third ray of dorsal
fin. Pelvic fin inserts slightly anterior to base ofpectoral fin. Fourth and first segmented rays
longest in pectoral and pelvic fins, respectively.
Fourth pelvic-fin segmented ray, if present,
minute and rudimentary. Caudal fin near truncate. Scales on body cycloid; head naked; lateralline scale row divided into two parts, i.e., anterior
and posterior series comprising grooved scales

New Creediine Fish from Ogasawara Islands

53

Fig. 3. Lateral view of head of Myopsaron nelsoni sp. nov., one of YCM-P 44207, 33.4 mm
SL. Solid and open arrows indicate slit of suborbital skin fold and a row of short cirri on
lower jaw, respectively.

and scales each with a distinct posterior lobe, respectively (Figs. 1, 4 and 5C).
Patterns of cephalic sensory canals are illustrated in Fig. 7. Oculoscapular canal with 7
pores; infraorbital canal interrupted below eye bythin skin fold, with 5 pores in anterior portionand 3 pores in posterior portion; preopercular
canal with 6–7 pores; mandibular canal well separated from preopercular canal, with 2 pores.
Osteology (Figs. 8, 9, 10A and 11–19). Cranium elongated, its greatest width ca. 2.6 in its
greatest length; interorbital area very narrow, its
least width ca. 4.7 in greatest width of cranium;
frontals with 2 pairs of large rounded orifices;
vomer toothless, with pointed anterior margin;
mesethmoid pointed anteriorly; lateral ethmoidwith broad lateral projection, bearing cartiliginous articular facet for palatine anterolaterally;
supraoccipital crest weakly developed; ventroanterior margin of basioccipital weakly notched. Infraorbital series completely encasing lower margin of orbit with 3 bones (including lachrymal)
and small fourth infraorbital (dermosphenotic);
second infraorbital tubular, much smaller thanlachrymal; third infraorbital slightly smaller thanlachrymal, with weakly developed posterior lobe.
Premaxilla toothless; articular process for maxilla and postmaxillary process of premaxilla welldeveloped, latter of which not indented dorsally;
ascending process autogenous, interlocked with

Fig. 4. Two types of lateral-line scales of Myopsaron nelsoni sp. nov., 1 of NSMT-P 94888, female, 32.5 mm SL. A) grooved lateral-line
scale in anterior series; B) pored lateral-line
scale with distinct posterior lobe in posterior
series.

main body of premaxilla anteriorly; ascending
process very long, more than 2 times articular
process in height; anterior end of ascending
process broadened, flat and upturned, supporting
cartilaginous tissue inside of globular extension
of upper jaw; rostral cartilage well developed, attached to tip of ascending process; maxilla with 3
heads, anterior and posteroventral ones over articular process of premaxilla, and remaining(dorsal) one connected with vomer via ligament(maxillo-vomerine ligament) with a reel-shaped
bony ossicle inside; posterior end of maxilla
forked, its ventral arm longer and more slenderthan dorsal arm; dentary bears 1–3 minute conical teeth just posterior to mid-dorsal margin;
coronoid process of dentary well developed, expanded dorsally and posteriorly, its dorsal tip

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Koichi Shibukawa

Fig. 5. Schematic illustration of three species of Creediinae, showing lateral-line scale rows (shaded area). A)
Limnichthys fasciatus, NSMT-P 34378, 50.0 mm SL; B) Creedia bilineata, based on illustration of holotype
by Shimada and Yoshino (1987, fig. 1) and 1 paratype examined (URM-P 6460, 24.5 mm SL); C) Myopsaron
nelsoni sp. nov., one of AMS I.44840-001, 34.3 mm SL.

Fig. 6. Lateral views of anterior parts of bodies
of Myopsaron nelsoni sp. nov., showing distinct sexual dimorphism in pectoral and pelvic
fins. A) male, 1 of AMS I.44840-001, 34.3 mm
SL; B) female, 1 of AMS I.44840-001, 36.3mm SL.

curved dorsoposteriorly; large, elongate ovoid
foramen at near mid-dorsal margin of dentary;
articular not or slightly overlapping dentary;palatine toothless, forked posteriorly, its dorsal
and ventral arms contact endopterygoid and ectopterygoid, respectively; endopterygoid largely
free except for anterior contact with palatine,
supporting ventral portion of eye socket; posterior tip of endopterygoid only slightly upturned,
but well apart from skull; ectopterygoid very
elongate, rod-like, with slightly broadened (butnot fan-shaped) posterior end attaching toquadrate and metapterygoid; preopercle, interopercle and subopercle splintered; distinct, large
open space circumscribed by margins of symplectic, hyomandibular, preopercle and quadrate(and symplectic well apart from preopercle). Ossified gill rakers developed on both sides of all
gill arches; gill rakers on outer surface of firstgill arch very low, small multispine laminae; remaining gill rakers raised as short stubs, fringed

New Creediine Fish from Ogasawara Islands

55

Fig. 7. Dorsal (top), lateral (middle) and ventral views (bottom) of head of Myopsaron nelsoni sp. nov. (one of
AMS I.44840-001, male, 35.0 mm SL), showing cephalic sensory canals (shaded area). AN and PN, anterior
and posterior nares, respectively.

by minute spines dorsally; basihyal long, rodlike; basibranchial 4 cartilaginous, elongate,
subequal to basibranchials 1 and 2 in length; epibranchials 1–4 simple, not forked medially;pharyngobranchial 1 absent; pharyngobranchials
2 and 3 connected via U-shaped cartilage anteriorly; pharyngobranchial 4 absent, fourth upper
pharyngeal tooth plate present. Urohyal notforked anteriorly; anterior tip of urohyal lies
under anterior end of basibranchial 2; anterior
five branchiostegal rays articulate with anterior
ceratohyal, and the remaining two with posterior

ceratohyal. Vertebrae 43–45, including 24–26
precaudal and 18–21 caudal (precaudal minus
caudal vertebrae 3–8, 7 in holotype); all neuraland haemal spines long, slender, not expanded
anteroposteriorly, excluding those of preural vertebra 2; neural spine of preural centrum 2 very
short, lying beneath anterior epural; haemal spineof preural centrum 2 slightly expanded anteroposteriorly; ribs absent; epineurals on all precaudal and anterior 3 caudal vertebrae; first dorsalfin pterygiophore inserts between 22nd and 23rd
(1 specimen), 23rd and 24th (11 specimens), or56

Koichi Shibukawa

Fig. 8. Dorsal (top) and ventral (bottom) views of neurocranium (skull) of Myopsaron nelsoni sp. nov., 1 of
NSMT-P 94888, female, 32.5 mm SL. Shading areas indicate cartilaginous parts.

24th and 25th (5 specimens, including holotype)neural spines; 1 (16 specimens, including holotype) or 2 (1 specimen) anal-fin pterygiophores
anterior to first haemal spine; middle radials of
dorsal- and anal-fin pterygiophores not present as
separate ossifications; distal radial of dorsal- andanal-fin pterygiophores cartilaginous posteriorly;
ultimate dorsal- and anal-fin pterygiophores lackdistal radial; hypurals and parhypural fused to
urostyle, forming urostylar complex; 2 epurals,
anterior one J-shaped. Cleithrum with 3 heads
dorsally, inner one of which directed posteriorly;
postcleithra absent; pelvis with dorsomediallyexpanded suprapelvic keel, forming an invertedbowl shape; anterolateral flange of pelvis (subpelvic keel) not or slightly expanded anteriorly;
iliac spur of pelvis minute, arising near pelvic
spine articulation and directed ventromedially;

postpelvic process undeveloped.
Color when fresh. No information available.
Color in alcohol (Fig. 2). Ground color ofbody pale yellow. Two dark brown stripes on
upper half of body; dorsal one along dorsal profile from nape to caudal fin base, ventral one on
lateral midline from a little behind base of pectoral fin to caudal-fin base; posterior margin ofeach scale pocket covered by these stripes slightly darkened. A short dark brown bar just above
pectoral-fin base, forming bridge between dorsal
and mid-lateral stripes on body. Several variously-shaped brown or dark brown markings on dorsal surface from snout to nape. A pupil-sized
dark brown spot on mid-lateral snout. Cheek
with 2 dark brown markings, comprising a small(smaller than pupil) dark brown spot below eye
and a short diagonal dark brown bar at mid-pos-

New Creediine Fish from Ogasawara Islands

57

Fig. 9. Lateral view of jaws, suspensorium and opercular bones of Myopsaron nelsoni sp. nov., 1 of NSMT-P
94888, female, 32.5 mm SL. Shading areas indicate cartilaginous parts, except for silhoutte of skull.
Table 1. Measurements of Myopsaron nelsoni gen. et sp. nov. Data for paratypes are shown in means, with ranges
in parentheses
Holotype

Paratypes

NSMT-P 94886

Males (n13)

Females (n3)Standard length (mm)35.6

31.8–36.3

32.5–36.3

In % of standard length
Head length
Snout length
Eye diameter
Interorbital width
Body depth
Predorsal length
Prepelvic length
Preanal length
Preanus length
Caudal-peduncle depth
Length of dorsal-fin base
Length of anal-fin base
Length of 1st ray of dorsal fin
Length of 2nd ray of dorsal fin
Length of longest ray of pectoral fin
Length of pelvic-fin spine
Length of 1st ray of pelvic fin
Length of 2nd ray of pelvic fin
Length of 3rd ray of pelvic fin
Length of 1st ray of anal fin
Length of 2nd ray of anal fin

22.1
6.2
4.2
1.4
9.1
64.3
20.2
66.4
62.8
4.5
29.5
33.0
9.6
10.4
23.0
5.8
25.8
10.8
6.2
5.6
8.2

22.7 (21.7–23.7)
6.7 ( 6.2–7.2)
4.3 ( 4.0–4.6)
1.5 ( 1.3–1.8)
8.9 ( 8.3–9.5)
63.3 (60.7–65.4)
21.5 (20.1–23.0)
65.3 (63.3–67.7)
62.7 (60.9–65.5)
4.5 ( 4.1–4.9)
28.6 (26.9–30.8)
32.1 (30.4–33.9)
9.2 ( 7.1–11.0)
10.6 ( 9.7–11.9)
22.4 (19.8–24.7)
5.6 ( 4.9–6.2)
25.0 (21.7–27.9)
9.4 ( 8.4–10.3)
6.1 ( 5.1–7.4)
5.3 ( 4.1–7.5)
7.6 ( 6.5–8.7)

23.3 (23.0–23.7)
6.9 ( 6.6–7.4)
4.5 ( 4.2–4.8)
1.4 ( 1.3–1.4)
9.2 ( 9.0–9.5)
64.7 (63.8–65.2)
21.3 (20.3–22.2)
66.6 (65.2–68.5)
63.8 (62.3–65.2)
4.4 ( 4.2–4.6)
27.3 (25.8–29.1)
30.0 (29.1–30.6)
9.4 ( 9.1–9.9)
9.9 ( 9.0–10.3)
9.1 ( 9.0–9.2)
2.3 ( 2.2–2.3)
4.4 ( 4.1–5.0)
3.1 ( 3.1–3.3)
2.2 ( 2.0–2.5)
5.0 ( 5.0–5.2)
6.9 ( 6.7–7.4)

terior part. A dark brown blotch just behind anterior tips of lower jaws. Dorsal, anal, pectoral andpelvic fins subtranslucent with pale rays. Caudal
fin with a dark brown transverse bar a little behind its base; sometimes an additional indistinct

brown transverse bar at middle of caudal fin.
Sexual dimorphism. Pectoral and pelvic finsare greatly elongate, long and filamentous inmale, whereas short and non-filamentous in female (Fig. 6; Table 1). Although similar sexual

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Koichi Shibukawa

dimorphism in these fins has been reported in 2
Creedia species (i.e., C. haswelli and C. partimsquamigera) by Nelson (1983), the degree of finelongation in male Myopsaron nelsoni is muchgreater than in Creedia. Within the type series ofM. nelsoni, the smallest male specimen withelongate fins is 22.1 mm SL (OMNH-P 35374).
The other known sexual dimorphism known increediine fishes, i.e. serrations on the posterior
lobe of the lateral-line scales (known only inmales of Creedia haswelli), is not present in
Myopsaron nelsoni.
Early development. One of the smallest specimens (16.6 mm SL) appears to be least developed within the specimens examined, as follows:eyes not typical creediine-like, i.e., lens not protruding beyond, and partially covered by, cornea;no spines along preopercular margin, but posteroventral corner of preopercle near angular(rather than rounded); all fin rays developed, ex-

cept pectoral- and pelvic-fin rays; scales undeveloped on body, except posterior lateral-line scales;
head, body and fins not pigmented, except eyeball black. In another 16.6 mm SL specimen, thelens protrudes beyond the cornea; the free rear
margin of the preopercle is rounded; all fin rays
are developed; both anterior and posterior lateralline scales are developed (but the other scales remain undeveloped). The smallest pigmented
specimen is 16.9 mm SL; in this specimen, a narrow area around the dorsal profile of the body isweakly pigmented by minute melanophores.
Along with increasing pigmentation and bodysize, the scaled area is radically broadened fromthe posterior part of the body in addition to thelateral-line scales (Fig. 20).
Reader and Neira (1998) listed “Small posterior preopercular spines from 3–5 mm, disappearing from 9 mm” as one of the main characters of
creediine (as creediid) larvae. All examined specimens of Myopsaron nelsoni (16.6–36.3 mm SL)
lack preopercular spines, but the spines might
have already disappeared in this size range.

Fig. 11. Lateral view of lower-jaw bones of Myopsaron nelsoni sp. nov., 1 of NSMT-P 94888,
female, 34.6 mm SL. Arrow indicates minutetooth on dentary.

Fig. 10. Dorsoanterolateral views of upper jawsof 2 species of Creediinae. A) Myopsaron nelsoni sp. nov., 1 of NSMT-P 94888, 32.5 mm
SL; B) Limnichthys fasciatus, MTUF-P 25670,
35.0 mm SL. Shading areas indicate cartilaginous parts, except for silhouettes of skulls.

Fig. 12. Lateral view of infraorbital bones of Myopsaron nelsoni sp. nov., 1 of NSMT-P 94888,
female, 32.5 mm SL. Note that the fourth infraorbital (or dermosphenotic) is not shown.

New Creediine Fish from Ogasawara Islands

Distribution and habitat. All specimens ofMyopsaron nelsoni were collected by sledge nets
from flat sandy bottoms at depths of 51–99 m off
the southern part of Chichi-jima Island, Ogasawara Islands.
Remarks. Some unusual extra ossicles andcartilaginous tissues are developed in the oral region of Myopsaron nelsoni (Figs. 9 and 10A).
These include: globular cartilaginous tissue witha shallow hollow ventrally, supported by the anterior end of ascending processes of the premaxillae; a reel-shaped bony ossicle in the maxillovomerine ligament; a small globular bony ossicle
in the premaxillo-rostral ligament; and a labial
cartilage (cartilaginous labial ligament). Of

Fig. 13. Dorsal view of branchial arches of Myopsaron nelsoni sp. nov., 1 of NSMT-P 94888,
female, 32.5 mm SL. Note that the dorsal elements of right side is removed, and shown in
Fig. 13. Shading areas indicate cartilaginousparts.

59

these, similar ossicles were illustrated by Shimada and Yoshino (1987, fig. 2) in Creedia bifasciata. Present research reveals that Limnichthys, theonly other creediine genus examined for its osteology, has a cartilaginous tissue within maxillovomerine ligament (Fig. 10B); this is, nevertheless, not hypertrophied and rather similar to themaxillo-vomerine meniscus found in most acanthomorphs (Johnson and Brothers, 1993). No
traces of the other extra ossicles and/or cartilaginous tissues were seen in Limnichthys. Nelson(1985) and Rosa (1995) did not refer to any ofthese extra ossicles and/or cartilaginous tissues intheir osteological surveys of the creediine fishes.
The extra ossicles and/or hypertrophied cartilaginous tissues around the oral region are notcommon but are known from several actinopterygians, e.g., gasterosteiform family Pegasidae,
atheriniform family Phallostethidae, and perciform families Epigonidae, Ammodytidae andSchindleriidae, with various degrees of development (Gosline, 1963; Ida, 1973; Pietsch, 1984;
Johnson and Brothers, 1993; Ida et al., 1994).
Presence of similar features in unrelated fishes
suggests that they represent examples of convergence correlated with jaw protrusion (Pietsch,1984).

Fig. 14. Ventral (left) and dorsal (right) views of
dorsal elements of left-side branchial arches ofMyopsaron nelsoni sp. nov., 1 of NSMT-P
94888, female, 32.5 mm SL. Shading areas indicate cartilaginous parts.

60

Koichi Shibukawa

Amongst the fishes with these hypertrophiedelements, the ammodytids are peculiar in having
autogenous ascending proceses of the premaxillae (Ida, 1973; Pietsch and Zabetian, 1990; Ida
et al., 1994) as do the trichonotids. The ammodytids also share another uncommon feature,a large anterolateral foramen in the dentary, withMyopsaron nelsoni and Creedia bilineata (Ida,

1973; Shimada and Yoshino, 1987; Ida et al.,
1994; present study). Pietsch and Zabetian(1990) indicated that the foramen found in
the ammodytids is “apparently unique among
teleosts.” Ammodytids are also known as sanddiving fishes, and the salient resemblances in aseries of uncommon features between ammodytids and creediines quicken our interest particularly in functional morphology. Note that thedentary foramen is apparently undeveloped in
Schizochirus, Limnichthys and the type species of
Creedia, viz. C. haswelli, regarding illustrations
or photographs of lower jaw of creediines shown
by Nelson (1985), Rosa (1995) and Smith andJohnson (2007); the condition in the other creedi-

Fig. 15. Lateral views of hyoid arches of Myopsaron nelsoni sp. nov., 1 of NSMT-P 94888,
fmale, 32.5 mm SL. Shading areas indicate
cartilaginous parts.

Fig. 17. Lateral view of caudal-fin skeleton of
Myopsaron nelsoni sp. nov., 1 of NSMT-P
94888, male, 34.6 mm SL. Shading areas indicate cartilaginous parts.

Fig. 16. Lateral views of axial skeleton and vertical fins of Myopsaron nelsoni sp. nov., 1 of NSMT-P 94888,
male, 34.6 mm SL.

New Creediine Fish from Ogasawara Islands

61

ines remains unexplored.
Etymology. The new species is named for J.
S. Nelson, in reference to his great contribution
to our knowledge about taxonomy and systematics of the creediine fishes.
Discussion
Nelson (1985) recognized monophyly in thecreediine (as creediid) clade on the basis of 21
characters. Myopsaron nelsoni agrees well withalmost all diagnostic features of the Creediinae
discussed by Nelson (1985), except as follows:
globular shaped fleshy extension at tip of upper
jaw, discontinuous lateral line on body, higher
number of precaudal than caudal vertebrae, andedentate premaxilla. These 4 characters arefound only in M. nelsoni within the family, andare regarded as autapomorphic for the species.
The other feature differing from what Nelson
showed is the position of “jaw articulation” (articular-quadrate articulation). As described by
Nelson (1985; 1986), the jaw articulation is positioned at the point posterior to vertical from theposterior margin of orbit in all creediine genera
except Myopsaron. However, the point of jaw articulation in Myopsaron is placed slightly forward to a vertical from the posterior margin of
the orbit (Fig. 9); it is more similar to that of thetrichonotine genus Trichonotus (see Nelson,
1986, fig. 1) than to that of other creediines. The
position of jaw articulation varies among thecreediines (see Nelson, 1985, figs. 2 and 3), and,considering the other osteological characteristics(see below), the condition in jaw articulation of
Myopsaron is hypothesized as the one extreme,
apparently plesiomorphic, example within the
Creediinae.
Another work defining the Creediinae was thatof Rosa (1995, as Creediidae). In the text of her
paper, she noted that the family is separated fromthe other trachinoids (sensu Pietsch, 1989) by
the following 3 “new” characters, suggesting its
monophyly: 1) dentary bone not completely
toothed (i.e., no teeth on symphysis and posterior
portion of dentary); 2) anterior tip of urohyal

62

Koichi Shibukawa

Fig. 20. Schematic illustrations of juveniles of Myopsaron nelsoni sp. nov., 3 specimens of NSMT-PL 723,
showing development of scaled areas. A) 17.0 mm SL; B) 17.5 mm SL; C) 19.2 mm SL.

lying on second or third basibranchial; 3) posterior tip of mesopterygoid reaching posterior margin of skull [sic.] (probably error of “orbit”). On
the other hand, in the abstract of the same paper,
she listed 4 characters as “new characters of taxonomic significance for the group”, including
above listed 1) and 2) and the other 2 characters,i.e., 4) lower arm of posttemporal not reachingskull, and 5) posterior tip of ectopterygoid reaching posterior margin of orbit; any accounts of thelast 2 features were not apparent in the text. Myopsaron possesses at least all of Rosa’s (1995)
features, exclusive of 3) and 5).
Interrelationships of the creediine genera were
explored by Nelson (1985), based on “similarities and differences” of 15 characters. He recognized 3 unresolved basal lineages within the subfamily; first and second lineages comprise
Schizochirus and Creedia, respectively, while
the third contains the remaining 5 genera(Apodocreedia, Chalixodytes, Crystallodytes,
Limnichthys and Tewara). Nelson (1985) assumed that Schizochirus is the most primitive
creediine genus, having, e.g., deeper body (comparing with the other creediines), branched raysof pectoral, pelvic and anal fins, wide inter-pelvic
distance, and appearance of lateral-line scales resembling some hemerocoetines. This assumption
is followed here. Multiple rows of suborbital- andpreopercular-canal pores, resembling that ofhemerocoetines, also suggest that Schizochirus is
the most primitive derivative within the family.
Myopsaron, Creedia and the other creediine genera have the specialized condition in these features, viz. body elongate, all pectoral, pelvic andanal-fin rays unbranched, inter-pelvic distancenarrower, lateral-line scales non-hemerocoetinelike, and suborbital- and preopercular-canalpores forming uniserial rows in each (Table 2).
All 8 creediine genera are compared in Table
2. Myopsaron is most similar to Creedia in sharing the following diagnostic, apparently derived,features within the family: 1) 12–16 dorsal-finrays; 2) relatively long, tube-like second infraorbital; 3) ventrally directed bony flange at anterior
margin of postocular broadened portion offrontals (see Rosa, 1995: 48, fig. 1). These 2 genera also share in having the suborbital skin folddiscontinuous midway (and anterior skin foldcovering ventral margin of eye), not found in thenot forked
short stubslightly upturned

undeveloped
forkeddeveloped

not forked
short stubslightly upturned

Type A
multiple
continuous
absentabsentpresent
present
present
2 pairs and single
medianundeveloped
forkeddeveloped

well developed

beak-likenot tubularabsentpresent or absent
well developed toreducedType B
uniserial
continuouspresent
absentpresent
present
present or absent
2 pairs and single
medianslightly expanded
forkeddeveloped
slightly to well
upturned
not forked
short stub

beak-like (but lessdeveloped)
not tubularpresent
present

forkedelongate

greatly upturned

Type A
uniserial
continuous
absentpresent
present
present
absent1 pair and single
medianexpanded
not forked
undeveloped

well developed

tubularabsentabsentbeak-like

21–33
25–34
10–16
4–5
8
37–45
14–16
37–45
1–2
52

5

Limnichthys

forkedelongate

greatly upturned

Type A?
?
absentpresent
present
present
absent1 pair and single
medianexpanded
not forked
undeveloped

well developed

tubularabsentabsentbeak-like

34–36
37–40
13–14
5
8
50–53
14
50–53
1
52

1

Tewara*1 Data is cited from Rosa (1995); Nelson (1985) gave a rage of 49–55 in vertabral count of Tewara cranwellae.
*2 Counts of branchiostegal rays are provided in the form of those attached to ceratohyalepihyal.
*3 Types of lateral line are illustrated in Fig. 5.

Anterior tip of urohyal (in dorsal view)
Neural spine of 2nd preural centra

Posterior end of endopterygoid

Posterior lobe of 3rd infraorbital
Posterior tip of maxilla
Postmaxillary process of premaxilla

2 pairs

Type C
uniserial
discontinuouspresent
absentabsentabsentabsentOrifices on frontals

well developed

Lateral-line arrangement*3
Pores of suborbital and preopercular canals
preopercular and mandibular canals
Slit-like extension on ventral margin of eye
Dorsal saddles on body
Teeth on vomer
Teeth on premaxilla
Teeth on palatine

not tubularabsentabsentScales on body

Anterior nostril
Branched anal-fin rays
Branched pectoral- and pelvic-fin rays

globular

Fleshy extension of upper jaw

12–16
17–28
10–15
3–4
8–9
40–47
14–16
41–47
2
52

4

Creedia

18–20
28–29
16–17
5
9
36–41
12
36–41
2
52

1

1

14–16
16–18
11–13
3–4
9
333–35
24–26
43–45
2
52

Number of species

Counts
Dorsal-fin rays
Anal-fin rays
Pectoral-fin rays
Pelvic-fin segmented rays
Branched caudal-fin rays
Lateral-line scales
Precaudal vertebrae
Total vertebrae
Epurals
Branchiostegal rays*2

Schizochirus

Myopsaron gen. nov.

not forked
elongate

greatly upturned

Type B?
?
absentabsentpresent or absent
present
absent2 pairs and single
medianexpanded
not forked
undeveloped

well developed

tubularabsentabsentbeak-like

35–40
32–36
13–14
5
8
55–58
23
55–58
1
61

1

Apodocreedia

forkedelongate

greatly upturned

Type A
uniserial
continuous
absentpresent
present
present
absent1 pair and single
medianexpanded
not forked
undevelopd

reducedtubularabsentabsentbeak-like

30–43
34–41
10–14
5
8
50–58
24-25
50–58*11
61

2

Crystallodytes

forkedelongate

greatly upturned

Type A
uniserial
continuous
absentpresent
present
present
absent1 pair and single
medianexpanded
not forked
undeveloped

reducedtubularabsentabsentbeak-like

36–40
36–40
12–13
4
8
56–59
23-24
56–59
1
61

2

Charixodytes

Table 2. Comparisons of the creediine genera. Data is cited from Nelson (1978, 1983, 1985), Nelson and Randall (1985), Shimada and Yoshino (1987), Rosa (1995)
and Yoshino et al. (1999), in addition to examination of the present study.

New Creediine Fish from Ogasawara Islands
63

64

Koichi Shibukawa

other creediines. Nevertheless, the phylogenetic
implication of this character is uncertain, because
a similar condition is found in the trichonotines(comprising Trichonotus only) and the othersand-diving fish family Ammodytidae. All ofthese fish groups with a discontinuous suborbitalskin fold are bottom-oriented elongate sanddivers, suggesting that the condition has been independently derived with their sand-diving habit.
Creedia and Myopsaron also share the following features, not found in other creediines except
Schizochirus: usually 9 branched caudal-fin rays
[except for intraspecific variation reported by
Nelson (1978) in C. haswelli]; posterior end of
maxilla forked; weakly to well developed postmaxillary process of premaxilla; free rear marginof preopercle with splintering; posteroventral
laminated projection of lacrymal undeveloped;
relatively long and slender urohyal; short neuralspine of second preural centrum (PU2) underepural(s). All of these features appear to be primitive within the Creediinae, since they are also
found in the other 2 trichonotid subfamilies, i.e.,
Trichonotinae and Hemerocoetinae (Nelson,
1986; Fujita, 1990; present study). Lack of specialized condition in these features suggests that,
as well as Schizochirus, Creedia and Myopsaron
do not belong to the lineage including the other 5genera.
A sister group relationship of Myopsaron with
Creedia is open to question. Although these 2
genera share some putative apomorphies as noted
above, Myopsaron appears to be a plesiomorphic
condition in the position of its jaw articulation
and a barely-developed posterior lobe of third infraorbital, which resembles the condition in mostprimitive derivate Schizochirus. Further study, including molecular analysis, is needed to resolve
certain relationships of the creediine genera.
Comparative materials. Chalixodytes
chameleontoculis: AMS I. 28101-010, 1 specimen, 41.9 mm SL, west of Riviere, Riviere desAnguilles, south coast at Senneville, Mauritius,
0.2–1.5 m depth, 1989 (collected by J. Paxton, O.
Griffiths and M. Welshman); AMS I. 41428-008,
5 specimens, 19.4–25.9 mm SL, Souillac, South-

ern Mauritius, Mauritius, Jan. 2002 (collected by
O. Griffiths). Chalixodytes tautensis: AMS I.
37908-010, 1 specimen, 41.1 mm SL, southeastern side of Efate Island, Vanuatu, 0–1 m depth, 6
May 1997. Creedia bilineata: URM-P 6460, 1
specimen (paratype of C. bifasciatus), 24.5 mm
SL, Hatoma-jima Island, Yaeyama Group of
Ryukyu Islands, Japan, 9 Nov. 1979 (collected by
K. Shimada); URM-P 6463, 1 specimen(paratype of C. bifasciatus: cleared, stained anddissected), Kabira Bay, Ishigaki-jima Island,
Yaeyama Group of Ryukyu Islands, Japan, 15
Oct. 1985 (collected by K. Shimada); URM-P
11750, 1 specimen, 36.6 mm SL. Creedia
haswelli: AMS B 9210-9213, 4 specimen (syntypes of Hemerocoetes haswelli), 44.3–50.7 mm
SL. Creedia partimsquamigera: AMS I. 21420001, 1 specimen (holotype of Creedia partimsquamigera), 53.8 mm SL, off Coogee Beach,
Sydney, New South Wales, Australia, 26 Jan.
1980 (collected by R. Kuiter); AMS I. 21420005, 2 specimens (paratypes of Creedia partimsquamigera), 57.5–66.1 mm SL, off Coogee
Beach, Sydney, New South Wales, Australia,
26 Jan. 1980 (collected by R. Kuiter); AMS I.
22868-001, 2 specimens (paratypes of Creedia
partimsquamigera), 38.3–55.9 mm SL, Sept.
1981 (collected by R. Kuiter). Crystallodytes
pauciradiatus: AMS I. 24606-001, 2 specimens(paratypes of Crystallodytes pauciradiatus),
30.3–31.3 mm SL, south end of Hanga Roa, offnorthern coast of Easter Island, 10 Feb. 1969
(collected by G. Allen and J. Randall).
Limnichthys fasciatus, 14 specimens (syntypes ofLimnichthys fasciatus), 21.3–36.1 mm SL, Lord
Howe Island, Feb. 1903 (collected by Waite andMcCulloch); MTUF-P 25670, 1 specimen(cleared and stained), Kominato, Chiba Prefecture, Japan, 29 Aug. 1984; NSMT-P 34378, 1
specimen, 50.0 mm SL, Koajiro, Miura City,
Kanagawa Prefecture, Japan, 26 Apr. 1991 (collected by M. Aizawa). Limnichthys polyradiatus:
AMS I. 19646-001, 1 specimen (paratype of
Limnichthys polyradiatus), 44.5 mm SL, near
Leigh, New Zealand, 18 Aug. 1971 (collected by
G. R. V. Anderson); AMS I.19647-001, 2 speci-

New Creediine Fish from Ogasawara Islands

mens (paratypes of Lymnichthys polyradiatus),
38.2–41.9 mm SL, Pink Beach, near Goat Island,
New Zealand, 1 Feb. 1972 (collected by G. R. V.
Anderson). Schizochirus insolens: AMS IA.
7062, 1 specimen, 34.9 mm SL, Ballina, NewSouth Wales, Australia (collected by M. Ward);
AMS IB 514, 1 specimen, ca. 52.2 mm SL (headdamaged), Mooloolabah, Queensland, Australia,
May 1940 (collected by T. Iredale); AMS I.
34384-010, 1 specimen, 21.5 mm SL, 2.5 km
western of Island Head, 0–6 m depth, 25 Oct.
1993 (collected by D. Bray and S. Reader).
Acknowledgments
I am grateful to T. Kanazawa (Oita Prefecture,
Japan) for providing all examined specimens ofMyopsaron nelsoni. I also wish to thank the following persons for their kind help during my visits to their institutions and/or registration of thetype series of M. nelsoni: M. McGrouther, D. F.
Hoese, J. Lewis, and A. C. Hay (AMS); H. Senou(KPM); H. Kohno (MTUF); K. Matsuura, G.
Shinohara and Y. Takata (NSMT); T. Yoshino
and T. Kon (URM: TK now belongs to Ocean
Research Institute, University of Tokyo); M.
Hayashi and K. Hagiwara (YCM). My sincere
thanks go to J. S. Nelson (University of Alberta,
Canada) for his critically reading a draft manuscript. I. L. Rosa (Universidade Federal da Paraiba CCEN Depto, Brasil) kindly provided literature. This study was supported in part by a research grant from the Fujiwara Natural History
Foundation, Japan.
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Manuscript received 7 August 2009; revised 1 February
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Associate editor: K. Matsuura.