Bull. Natl. Mus. Nat. Sci., Ser. A, Suppl. 4, pp. 89–95, March 29, 2010

Hetereleotris exilis, a New Goby (Teleostei, Perciformes, Gobiidae) fromthe Ryukyu 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 species of the gobiid fish, Hetereleotris exilis, is described based on a single
female specimen (22.1 mm SL) collected at the depth of 53 m of off Nagan’nu Island, Okinawa
Group of the Ryukyu Islands, Japan. It is unique within the genus in having very slender body
(depth at anal-fin origin 11.9% of standard length), and characterized by the following: VI-I, 12
dorsal-fin rays; I, 11 anal-fin rays; 16 pectoral-fin rays; I, 5 pelvic-fin rays; innermost (fifth)
segmented pelvic-fin ray unbranched, long, subequal to preceding ray in length; scales cycloid,
restricted on posterior part of body (ca. 14–15 longitudinal scales); no head sensory-canal pores.
Key words : Hetereleotris exilis, New species, Gobiidae, Gobiinae, Japan.

Gobies of the genus Hetereleotris Bleeker,
1874 are small-sized (up to ca. 50 mm SL) bottom dwellers with cryptic habit, typically foundin shallow coastal marine waters in the IndoWest Pacific. Hetereleotris belongs to the gobiid
subfamily Gobiinae (sensu Pezold, 1993), and isdistinguished from the other gobiine genera inhaving the following combination of characters(Akihito and Meguro, 1981; Hoese, 1986): firstgill slit closed by membrane from inner surfaceof gill cover to half or more of lower limb of firstgill arch; pelvic fins entirely separated one another, without frenum and connecting membrane between innermost rays [except for a single species
Hetereotris zanzibarensis (Smith, 1958), which
often has pelvic fins fused into a cup-shaped
disc]; posterior naris with an elevated rim or
opening at tip of a short tube; a single-lobed
mental frenum, followed by a pair of short longitudinal rows of sensory papillae; transverse pattern of sensory papillae on cheek; 101727
vertebrae; dorsal-fin pterygiophore formula (“PV” of Akihito, 1984) 3/II II I I 0/9.
Hoese and Larson (2005) proposed that Pascua Randall, 2005, described for a single species
Pascua caudilinea Randall, 2005 from the Easter
Islands, was a junior synonym of Hetereleotris.

One-year later, Randall (2006) refuted the Hoeseand Larson’s suggestion, and recognized 3
species within Pascua, viz. P. caudilinea, Pascua
readerae (Hoese and Larson, 2005) and Pascua
sticta (Hoese and Larson, 2005), latter 2 ofwhich have been described as the species of Hetereleotris by Hoese and Larson (2005) from theSouth Pacific. These 3 Pascua-type species sharemany features with the typical Hetereleotris, but
are readily distinguished from the latter by having: modified basicaudal scales with enlarged
cteni; flattened and elongate urogenital papilla ofmales; simple pore-like posterior naris (or with
only a slightly elevated margin anteriorly); andmuch reduced sensory-papillae rows on head
(Hoese and Larson, 2005; Randall, 2006). All ofthese features exclusive of urogenital papilla are,nevertheless, appeared as intra-generic variations
in some Indo-Pacific gobiin genera; namely, similar modification of basicaudal scales is found insome species of Cabillus Smith, 1959 (seeGreenfield and Randall, 2004), and similar intrageneric variations of sensory-papillae arrangement and condition of posterior naris are foundin Priolepis Valenciennes in Cuvier and Valenciennes, 1837 and Trimma Jordan and Seale, 1906
(see Winterbottom and Burrige, 1992, 1993). On

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Koichi Shibukawa

the other hand, as pointed out by Hoese and Larson (2005), shape of male urogenital papilla ofPascua resembles a part of Eviota Jenkins, 1903
(rather than typical Hetereleotris). Basic configuration of sensory-papillae rows on cheek in Pascua, e.g., sensory papillae row b short and close
to preopercular margin, also resembles that of
Eviota (vs. row b well apart from preopercularmargin in Hetereleotris). Further analysis on theinterrelationships between species of Hetereleotris, Pascua and the other gobiine genera
including Eviota is needed to evaluate the validity of Pascua, but that work is beyond the scopeof this paper.
Following the Randall’s (2006) concept ofgeneric limit provisionally, Hetereleotris comprises 14 species (Akihito and Meguro, 1981; Hoese,
1986; Gill, 1998) in addition to the new species
described here: Hetereleotris apora (Hoese andWinterbottom, 1979); Hetereleotris bipunctata
Tortonese, 1976; Hetereleotris caminata (Smith,1958); Hetereleotris diademata (Rüppell, 1830);
Hetereleotris georgegilli Gill, 1998; H. kenyae
Smith, 1958; Hetereleotris margaretae Hoese,
1986; Hetereleotris nebulofasciata (Smith,1958); Hetereleotris poecila (Fowler, 1946); Hetereleotris tentaculata (Smith, 1958); Hetereleotris vinsoni Hoese, 1986; Hetereleotris vulgaris (Kluzinger, 1871); H. zanzibarensis; Hetereleotris zonata (Fowler, 1934).
During the biological survey in the Ryukyu
Archipelago made by R/V Toyoshio-maru on
May 2004, a single specimen of unnamed speciesof Hetereleotris was dredged at the depth of 53 mof off Nagan’nu Island, Okinawa Group of
Ryukyu Islands, Japan. The species, herein described as new, is readily distinguished from thecongeners by the combination of cephalic sensory systems, meristic counts, shape of body, squamation, and coloration.
Materials and Methods
The specimen examined is deposited in thefish collection of the Department of Zoology, National Museum of Nature and Science, Tokyo(NSMT-P).
All fish lengths given are standard lengths(SL). Measurements were made point-to-pointwith calipers under the dissecting microscope tothe nearest 0.01 mm. The methods for measurements followed those of Hubbs and Lagler(1958), with exceptions given below (the snout
tip refers to the mid-anteriormost point of theupper lip): interorbital width was the least widthof bony interorbital space; jaw length was measured between the snout tip and the posteriormost
point of lip; head width and depth were measured
at preopercular margin; body depth was measured in 2 ways, the first at the first dorsal-fin origin, and the second at the anal-fin origin; napewidth was measured between dorsalmost margins
of gill openings; preanal and prepelvic lengths
were measured from the snout tip to the origin ofeach fin; pectoral-fin length was measured fromthe base to the tip of the longest ray; pelvic-fin
length was measured between the base of pelvicfin spine and the distal tip of the longest segmented ray; caudal-fin length was measured fromthe base to the tip of the middle caudal-fin ray.
The methods of counts followed Akihito (1984),
except for the following: gill rakers including all
rudiments were counted on the outer side of first
arch; count of pseudobranchial filaments included all rudiments. In order to confirm scales on
body, the holotype was stained by Alizarin Red(after taking photographs for making Fig. 1).
Scales and paired-fin rays were counted on bothsides. Osteological features were observed fromradiographs. The methods of Akihito (1984)
were used in describing the pattern of the interdigitation of the dorsal-fin pterygiophores between the neural spines (“P-V”). Cephalic sensory papillae were observed on the specimen
stained with cyanine blue, and notations on themfollow Miller (1986). All photographs and illustrations were made by the author.

New Species of Hetereleotris from Japan91

Fig. 1. Dorsal (top), lateral (middle) and ventral (bottom) views of Hetereleotris exilis sp. nov., NSMT-P 94885,
holotype, female, 22.1 mm SL, off Nagan’nu Island, Okinawa Group of Ryukyu Islands, Japan.

Hetereleotris exilis sp. nov.
[New Japanese name: Sasagaki-haze](Figs. 1–3)

Holotype. NSMT-P 94885, female, 22.1 mm SL, off
Nagan’nu Island, Okinawa Group of the Ryukyu Islands,
Japan (26°14.63N, 127°31.78E), 53 m depth, 22 May
2004, R/V Toyoshio-maru.

Diagnosis. Hetereleotris exilis differs from thecongeners in having the following combinationof characters: VI-I, 12 dorsal-fin rays; I, 11 analfin rays; I, 5 pelvic-fin rays; innermost (fifth)
segmented pelvic-fin ray unbranched, long, subequal to preceding ray in length; body elongate,
body depth at anal-fin origin 11.9% of SL; scales
cycloid, restricted on posterior part of body (ca.
14–15 longitudinal scales); no sensory canals
and associated pores on head.
Description. Dorsal-fin rays VI-I, 12; anal-fin

rays I, 11; pectoral-fin rays 16/16; pelvic-fin rays
I, 5/I, 5; segmented caudal-fin rays 98, including 87 branched rays; dorsal unsegmented caudal-fin rays 3; ventral unsegmented caudal-fin
rays 4; longitudinal scales approximately 14/15;
predorsal scales 0; gill rakers 20; vertebrae
1017=27; P-V 3/II II I I 0/9; epural 1; anal-fin
pterygiophores anterior to first haemal spine 2.
The following measurements are % of SL:
head length 27.1; head width 15.4; head depth
12.7; snout length 6.0; eye diameter 6.2; interorbital width 1.1; nape width 12.5; jaw length 10.1;
body depth at origin of first dorsal fin 14.7; body
depth at origin of anal fin 11.9; body width 14.4;
predorsal length 35.5; prepelvic length 26.7; preanal length 57.3; caudal-peduncle length 16.4;
caudal-peduncle depth 8.9; length of first dorsalfin base 18.2; length of second dorsal-fin base33.1; length of anal-fin base 28.7; pectoral-fin
length 22.7; pelvic-fin length 20.2; length of first

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Koichi Shibukawa

spine of first dorsal fin 10.6; length of second
spine of first dorsal fin 11.8; length of third spineof first dorsal fin 12.2; length of fourth spine offirst dorsal fin 11.4; length of spine of second
dorsal fin 11.0; length of first segmented ray of
second dorsal fin 14.0; length of longest (ninth)segmented ray of second dorsal fin 16.2; lengthof anal-fin spine 7.9; length of first segmented
ray of anal fin 11.3; length of longest (tenth)segmented ray of anal fin 15.0; length of pelvicfin spine 5.9; length of first segmented ray ofpelvic fin 9.8; length of fourth segmented ray ofpelvic fin 19.0; length of fifth segmented ray ofpelvic fin 18.7; caudal-fin length 24.3.
Body elongate and compressed. Head depressed, its depth 82.0% of width. Snout short,
its length subequal and slightly shorter than eye
diameter; snout does not protrude beyond upperlip. Eye dorsolateral, relatively small, its diameter 23.6% of head length; interorbital space narrow, its width narrower than pupil diameter and4.1% of head length. No cutaneous ridge along
dorsal midline of nape. Gape oblique, forming anangle of about 35 degrees with body axis. Lower
jaw slightly projecting beyond upper jaw; posterior end of jaws reaching to below middle (leftside) or anterior margin (right side) of pupil; jaw
length 37.1% of head length. Anterior and posterior nasal nares at tip of a short tube in each; nofleshy flap at tip of narial tubes; posterior narial
tube closer to eye than anterior narial tube. Tip of
tongue near rounded and very weak trilobate,
free from floor of mouth. Posteroventral marginof lower lip interrupted at symphysis. Mental
frenum weakly developed, forming small bump
just behind lower-jaw synphysis. Gill opening
narrow, restricted on base of pectoral fin. No
fleshy projections on lateral wing of shoulder girdle. No bony projections along posterior marginof preopercle. First gill slit largely closed bymembrane, i.e., ventral arm of first gill arch entirely continuous to inner surface of hyoid arch
via thin membrane; gill rakers on outer surface offirst gill arch greatly reduced, only 2 minutepapilla-like rakers on dorsal arm. Caudal peduncle moderately slender, its depth 54.0% of cau-dal-peduncle length. First dorsal fin slightly
lower than second dorsal fin; first dorsal fin close
to, but not connected to, second dorsal fin bymembrane; third spine of first dorsal fin longest(103.0% of preceding spine in length); first dorsal fin just reaching posteriorly to base of spineof second dorsal fin when adpressed; all dorsalfin spines slender and flexible, but not filamentous; all segmented rays of second dorsal fin
branched; ninth segmented ray of second dorsal
fin longest; second dorsal fin just reaching to avertical through caudal-fin base when adpressed.
Origin of anal fin slightly behind a vertical withbase of first segmented ray of second dorsal fin;height of anal fin slightly lower than second dorsal fin; anal-fin spine slender and flexible; all segmented anal-fin rays branched; tenth segmented
ray of anal fin longest; anal fin not reaching to avertical through caudal-fin base when adpressed.
Caudal fin rounded, almost symmetrical dorsoventrally; caudal fin slightly shorter than head, i.e.,
its length 89.6% of head length. Pectoral fin nearlanceolate, not reaching posteriorly to a verticalline through posterior end of base of first dorsal
fin or anus; all pectoral-fin rays branched. Originof pelvic fin far anterior to a vertical line throughorigin of first dorsal fin; pelvic fins entirely separated one another, without frenum and connected
membrane between innermost segmented rays;
all segmented pelvic-fin rays branched, except for
innermost (fifth) ray unbranched; fourth andfifth segmented rays subequal in length; pelvic
fin not reaching posteriorly to anus when adpressed. Urogenital papilla short and rounded.
All scales cycloid; scaled area restricted on
posterior part of body (Fig. 2), extending forward
to a vertical lines through interspaces between
bases of fifth and sixth (left side) or third andfourth (right side) segmented rays of second dorsal fin; anterior few scales non-imbricate andslightly embedded; scales on mid-lateral part ofposterior half of caudal peduncle largest, subequal to pupil in size; head and the other part ofbody naked.
All teeth on jaws unicuspid; upper jaw with 4
rows of teeth anteriorly, narrowing to single row

New Species of Hetereleotris from Japan

93

Fig. 2. Schematic illustration of lateral view of Hetereleotris exilis sp. nov. (NSMT-P 94885, holotype, female,
22.1 mm SL), showing scaled area (shaded) and pigmented pattern on head and body. Pectoral fin and segmented rays of all other fins are omitted. Note that many of scales were rubbed off, and recovered based on
the scale pockets in this drawing.

posteriorly; teeth on anterolateral part of outermost row largest, distinctly larger than teeth on
inner rows; lower jaw with 4 rows of teeth anteriorly, narrowing to single row posteriorly; teeth onanterolateral part of outermost row largest; noenlarged, distinct canine-like teeth on jaws; noteeth on vomer or palatine.
Cephalic sensory systems are illustrated inFig. 3. No sensory canals and associated pores onhead. Distinct (but, comparing the congeners,
somewhat reduced) transverse pattern of sensory
papillae rows on cheek; 4 transverse rows of sensory papillae below eye, posteriormost one(row 4) longest; row b short, close to row 4;
row d reduced and broadly interrupted between
end of jaws and mid-cheek, its posterior partcomprising 2 papillae; rows e and i uniserial (i.e.,
not containing transverse rows) in each; a pair ofshort longitudinal rows of sensory papillae justbehind chin (row f).
Coloration. Coloration when alive or fresh is
unknown. Color in 70% alcohol is as follows:
ground color of head and body pale yellow; 2radiated dusky bars from ventral margin of eye,viz., one between eye and posterior part of upper
jaw, and the other between eye and middle ofcheek; a patch of melanophores (about size ofeye) mid-lateral body below anterior part of baseof first dorsal fin; a mid-lateral series of ca. 5
faint minute patches of melanophores (comprising some melanophores in each) on tail, the lastof which located at end of hypural plates (that ofleft side almost faded); 4 saddle-like patches ofmelanophores on occipital region and nape; 2

Fig. 3. Schematic illustration of ventral (top),
lateral (middle) and dorsal (bottom) views ofhead of Hetereleotris exilis sp. nov. (NSMT-P
94885, holotype, female, 22.1 mm SL), showing cephalic sensory papillae (dots). AN and
PN, anterior and posterior nares, respectively.
Arrows show position where gill membrane is
attached to body. Note that the sensory-papillae row 3 on left side of this specimen appears
to be missing, and, in the middle figure, recovered the condition based on that on right side.

and 6 small saddle-like dusky spots along basesof first and second dorsal fins, respectively; 2faint, small saddle-like dusky blotches on caudal
peduncle; dorsal fins almost transparent, except
for single to several basal melanophores justabove saddle-like blotches along dorsal-fin bases;
anal, caudal and pelvic fins transparent; pectoral

94

Koichi Shibukawa

fin almost transparent, except for single (rightside) or 2 (left side) melanophores at around baseof fifth of sixth ray.
Distribution and habitat. The new species
Hetereleotris exilis is known only by the holotype, dredged at a depth of 53 m of off Nagan’nu
Island, Okinawa Group of the Ryukyu Islands,
Japan. No habitat data is available.
Comparison. The new species Hetereleotris
exilis clearly belongs to Heterelereotris, having
all diagnostic features listed above. Within thegenus, H. exilis is unique by having its elongate
body, i.e., body depth at anal-fin origin 11.9% of
SL vs. 13.9% or more of SL in the other congeners (Smith, 1958; Hoese and Winterbottom,
1979; Hoese, 1986; Gill, 1998).
Four species of Hetereleotris are known lacking head sensory canals and associated pores in
addition to Hetereleotris exilis, viz. H. apora, H.
bipunctata, H. diademata and H. margaretae. Of
these, H. apora has ctenoid scales on caudal peduncle (vs. all scales cycloid in H. exilis), with
4–5 longitudinal scales (vs. ca. 14–15); additionally, H. apora differs from H. exilis in having
10–11 segmented dorsal-fin rays (vs. 12), 9–10
anal-fin rays (vs. 11), no or a rudimentary fifth
segmented ray of pelvic fin (vs. fifth segmented
ray of pelvic fin well developed), 2 posteriorlydirected pointed spines on posterior margin of
opercle (vs. no such spines). Hetereleotris
bipunctata has 40–52 longitudinal scales (vs. ca.
14–15), usually 13 segmented dorsal-fin rays (vs.
12), usually 12 anal-fin rays (vs. 11), 15 pectoralfin rays (vs. 16), and a large humeral black spot
(vs. humeral spot absent). Hetereleotris diadema
lacks scales on body (vs. scales present), and has
a dark spot or faint band at end of caudal peduncle (vs. no dark markings at end of caudal peduncle), a dark spot dorsally at end of caudal peduncle (vs. absent), and 4–5 transverse papillae rows
under lower jaw (vs. no transverse papillae rows
under lower jaw). Hetereleotris margaretae has
28–30 longitudinal scales (vs. ca. 14–15), 11 segmented dorsal-fin rays (vs. 12), 10 segmented
anal-fin rays (vs. 11), 17 pectoral-fin rays (vs.
16), a pupil-sized dark brown spot above posteri-or end of operculum (vs. absent), and banded
pattern of body (vs. absent). All these data on
species of Hetereleotris referred to above are from
Hoese (1986), exclusive of those on H. exilis.
Other than Hetereleotris exilis, only a single
species, H. poecila, is known from the West Pacific. Hetereleotris poecila is a densely-pigmented, relatively deep bodied species (vs. body slender and very weakly pigmented in H. exilis), and,also, easily distinguished from H. exilis by thefollowing features (Akihito and Meguro, 1981;
Hoese, 1986): 9–10 segmented dorsal-fin rays
(vs. 12 in H. exilis); 8–9 segmented anal-fin rays
(vs. 11); body broadly scaled, 32–35 longitudinalscales (vs. scales restricted on posterior part ofbody, ca. 14–15 longitudinal scales); uppermost
2 rays of pectoral fin largely free from fin membrane (vs. no free pectoral-fin rays); cephalic sensory canal with pores B, C (unpaired), D (unpaired), E, F and H (vs. sensory canals and associated pores absent). Hetereleotris poecila is alittoral species as in many other congeners, foundin very shallow rocky shore at the depths of
0.2–5.0 m (Suzuki et al., 2009).
Remarks. The new species Hetereleotris exilis is known only from the holotype, dredged at
53 m depth. The other congeners have been collected from much shallower waters at the depthsof 0–12 m (Hoese, 1986; Gill, 1998; Suzuki
et al., 2009).
The holotype of Hetereleotris exilis is mature
female, with numerous developed eggs in theovary.
Etymology. The new specific name, exilis (aLatin word, meaning “slender”), refers to its slender body. The Japanese vernacular is a combination of the Japanese words “Sasagaki”, indicating
the finely-sliced pieces of vegetables (e.g., burdock), commonly used in the Japanese cuisines,and “Haze”, meaning “a goby”, in reference toits slender body.
Acknowledgments
I express my sincere thanks to H. Saitoh(NSMT) for providing the holotype of Hetereleotris exilis; G. Shinohara (NSMT) helped inNew Species of Hetereleotris from Japan

registration of the specimen; T. Suzuki (Hyogo
Prefecture, Japan) kindly provided literature; D.
F. Hoese (Australian Museum) read critically adraft manuscript.
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Manuscript received 22 August 2009; revised; accepted
17 February 2010.
Associate editor: S. Kimura