Bull. Natl. Mus. Nat. Sci., Ser. A, Suppl. 1, pp. 123–136, March 22, 2007

Grallenia, a New Goby Genus from the Western Pacific, with
Descriptions of Two New Species (Perciformes: Gobiidae: Gobiinae)
Koichi Shibukawa1 and Akihisa Iwata2
1

Department of Zoology, National Museum of Nature and Science, 3–23–1
Hyakunin-cho, Shinjuku-ku, Tokyo 169–0073, Japan
E-mail: shibu@kahaku.go.jp
2
Graduate School of Asian and African Area Studies, Kyoto University, 46
Shimoadachi-cho, Yoshida, Sakyo-ku, Kyoto 606–8501, Japan
E-mail: iwata@asafas.kyoto-u.ac.jp

Abstract A new gobiid genus, Grallenia, is described based on 2 new species from the western
Pacific. Grallenia belongs to the gobiid subfamily Gobiinae, and is distinguished from the othergobiine genera in having the following combination of characters: usually VI-I, 7–10 dorsal-finrays; I, 7–9 anal-fin rays; segmented caudal-fin rays 787, most frequently 87; gill opening
narrow, restricted (or nearly so) to pectoral base; first gill slit moderately open for most of its
length, but only slightly closed by membrane ventrally; anterior margin of tongue deeply concave;
pelvic fins fused medially together by well developed interradial membrane; pelvic frenum low orabsent; no distinct fleshy lobes developed around pelvic-fin spine; sensory-papillae rows on cheekrelatively reduced, with no distinct transverse rows; cephalic sensory canals present [with pores C(unpaired), D (unpaired), E and F] or absent; 1017182728 vertebrae; typical P-V patternof 3/II II I I 0/9; a pair of diagonal, elongate black spots at dorsal and ventral corners of posteriormost part of caudal peduncle. The new species Grallenia arenicola (13 types and 4 non-type specimens, 9.422.6 mm SL, Kashiwa-jima Island, Kochi Prefecture, Japan) differs from the congener
G. lipi (9 types and 2 non-type specimens, 10.8–18.8 mm SL, Ambon Island, Indonesia) in having:
second dorsal-fin rays I, 7–8, usually I, 7 (vs. I, 9–10 in G. lipi); anal-fin rays I, 7–8, usually I, 7
(vs. I, 9); vertebrae 1017182728, usually 101727 (vs. 1017182728, usually
101828); dorsal-fin spines not elongated in either sex (vs. first spine of first dorsal-fin greatly
elongated, filamentous in males); second dorsal fin not reaching caudal-fin base when adpressed,
length of longest ray 14.0–17.6% of SL (vs. usually reaching to, or beyond, caudal-fin base when
adpressed in adult, length of longest ray 17.5–21.8% of SL); length of longest ray of anal fin
8.9–13.3% of SL (vs. 12.6–18.1% of SL); scales on body extending anteriorly to just behind pectoral-fin base (vs. scales on body restricted on posterior half of body, except for isolated patch ofscales just behind pectoral-fin base); weak, low pelvic frenum (vs. pelvic frenum absent); length ofpectoral fin 20.7–25.9% of SL (vs. 23.2–30.6% of SL); caudal-peduncle length 24.6–28.6% of SL (vs.
20.3–23.6% of SL); cephalic sensory canals with pores C (unpaired), D (unpaired), E and F (vs. no
cephalic sensory canals and associated pores); no orange line on dorsal fins (vs. orange line at middleof dorsal fins in life). Fishes of Grallenia are small-sized (less than 23 mm SL) sandy-bottom dwellers
found on flat, fine and clean sandy bottoms, not commensal with any marine organisms.
Key words : Grallenia, Gobiidae, new genus, new species, western Pacific.

In the present paper, we describe a new marine
goby genus, based on 2 new species from thewestern Pacific. The species of this genus, belonging to the gobiid subfamily Gobiinae sensu
Pezold (1993), are diminutive (less than 23 mm

in standard length), slender subtranslucent bottom dwellers, found on flat, fine and clean sandy
bottoms around rocky/coral reefs (to 50 m depth).
They were already reported as unidentified gobies in some recently-published pictorial books of

124

K. Shibukawa and A. Iwata

marine fishes (Kuiter and Tonozuka, 2001; Allenet al., 2003; Senou et al., 2004).
Materials and Methods
Institutional abbreviations follow Leviton et al.
(1985), except for KPM (Kanagawa PrefectualMuseum of Natural History, Japan). Species accounts are presented in alphabetical order.
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 width
between innermost rims of right and left eyeballs; jaw length was measured between thesnout tip and the posteriormost point of lip; headwidth and depth were measured at preopercularmargin; body depth was measured at the anal-fin
origin; head depth and width were measured at
preopercular margin; nape width was measured
between dorsalmost margins of gill openings;
preanal and prepelvic lengths were measured
from the snout tip to the origin of each fin; pectoral-fin length was measured from the base to
the tip of the longest ray; pelvic-fin length wasmeasured between the base of pelvic-fin spine
and the distal tip of the longest segmented ray;caudal-fin length was measured from the base to
the tip of the middle caudal-fin ray. The methodsof counting followed Akihito (1984), except for
the following: longitudinal scale count was thenumber of oblique (anterodorsal to posteroventral) rows starting from and including the anteriormost scale and proceeding posteriorly to the
mid-base of caudal fin (scaled area of Grallenia
lipi is divided into two parts, and, thus, the longitudinal scales in anterior and posterior scaled
areas were counted separately); three methods oftransverse scale counts were taken (see descriptive accounts); circumpeduncular scale count was
the number of zigzag rows along a vertical line
around the caudal peduncle (counted from thesecond middorsal scale prior to the anteriormost
upper procurrent caudal-fin ray); gill rakers including all rudiments were counted on the outerside of first arch; count of pseudobranchial filaments included all rudiments. Scales (except for
predorsal and circumpeduncular scales) andpaired-fin rays were counted on both sides, but
gill rakers and pseudobranchial filaments arecounted on the right side only. Osteological features were observed from radiographs (for all
specimens) and some cleared and stained specimens were prepared following the methods ofPotthoff (1984). 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 canals and papillae were observed on specimens stained with cyanine blue, and notations onthem followed Akihito (1984) and Miller (1986),
respectively.
Descriptions of species were based only on thetype specimens; in describing meristic counts ofeach species, the count of the holotype is asterisked, and the number in parentheses after thecount indicates the number of specimens withthat count.
Grallenia gen. nov.
(New Japanese name: Sazare-haze zoku)

Type species. Grallenia arenicola Shibukawa
and Iwata, new species.
Included species. Grallenia comprises 2
species, G. arenicola and G. lipi, both of which
are described herein as new. See also “Remarks”
of G. lipi.
Diagnosis. Grallenia belongs to the gobiid
subfamily Gobiinae (sensu Pezold, 1993), and isdistinguished from the other gobiine genera inhaving the following combination of characters:
usually VI-I, 7–10 dorsal-fin rays; I, 7–9 anal-finrays; all dorsal- and anal-fin spines slender andflexible; segmented caudal-fin rays 787 (almost always 87); gill opening narrow, restricted

New Goby Genus from Western Pacific(or nearly so) to pectoral base; first gill slit moderately open for most of its length, but only
slightly closed by membrane ventrally; anterior
margin of tongue deeply concave; pelvic fins
fused medially by well developed interradial connecting membrane; pelvic frenum low (G. arenicola) or absent (G. lipi); no distinct fleshy lobes
developed around pelvic-fin spine; sensory-papillae rows on cheek relatively reduced, with no distinct transverse rows and row e not or just extending around posterior corner of jaws; cephalicsensory canals present [G. arenicola, with poresC (unpaired), D (unpaired), E and F] or absent(G. lipi); 101718=2728 vertebrae; typicalP–V pattern of 3/II II I I 0/9; 2 anal-fin pterygiophores anterior to first haemal spine; a pair of diagonal, elongate black markings at dorsal andventral corner of posteriormost of caudal peduncle. Fishes of Grallenia are small-sized (lessthan 23 mm SL) sandy-bottom dwellers, notcommensal with any marine organisms.
Description. Dorsal-fin rays VI–VII-I, 7–10;
anal-fin rays I, 7–9; pectoral-fin rays 14–17 (usually 15 or 16); pelvic-fin rays I, 5; segmented
caudal-fin rays 787 (most frequently 87),including 575 branched rays; dorsal unsegmented caudal-fin rays 5–9; ventral unsegmented
caudal-fin rays 5–8; longitudinal scale rows 10–
29; transverse scale rows from anal-fin origin
dorsoanteriorly to base of first dorsal fin 4–7 (inG. arenicola) or 0 (in G. lipi); transverse scalerows from anal-fin origin dorsoposteriorly tobase of second dorsal fin 6–9 (in G. arenicola) or
0 (in G. lipi); transverse scale rows from second
dorsal-fin origin ventroposteriorly to base of anal
fin 6–8 (in G. arenicola) or 0 (in G. lipi); predorsal scales 0; circumpeduncular scales 12; gillrakers on outer surface of first arch 033
9412; pseudobranchial filaments 3–6.
Body moderately elongate, subcylindrical,
compressed posteriorly. Head subcylindrical orslightly depressed, its depth 84.8–97.1% ofwidth. Snout pointed, subequal or shorter thaneye diameter, snout length 47.8–101.3% of eye
diameter. Eye dorsolateral, large, its diameter
26.9–36.6% in head length; eyes close together

125

dorsomedially, interorbital width narrower thanpupil diameter. No raised cutaneous ridges, flaps
or barbels on head. Mouth subterminal, slightly
oblique, forming an angle of about 35–40 degrees with body axis; lower jaw slightly to wellprojecting beyond upper jaw; posterior end ofjaws extending between vertical lines through anterior margin of eye and middle of pupil. Anterior naris opening at tip of short tube without skinflaps; distal tip of anterior narial tube not reaching to upper jaw when adpressed; posterior narial
opening with low rim, located about midway between anterior naris and eye. Anterior margin of
tongue deeply concave; tip of tongue free fromfloor of mouth. Ventral free edge of lower lip interrupted at symphysis; small bulbous mental
protrusion present or absent just behind chin.
Gill opening narrow, restricted (or nearly so) topectoral-fin base. No fleshy papillae on lateral
margin of lateral wing of cleithrum. No bonyprojections along posterior margin of preopercle.
Dorsal fins well separated from one another; firstdorsal fin subequal or slightly higher than second
dorsal fin, excepting males of G. lipi that have agreatly elongated, filamentous first spine; in
adults, all segmented dorsal-fin rays branched except anterior 1–2 rays. Anal fin slightly lower
than second dorsal fin in height; anal fin originates at vertical through base of spine or first
segmented ray of second dorsal fin; in adults, allsegmented anal-fin rays branched except anterior
1–2 rays. Caudal fin almost symmetrical dorsoventrally, nearly rounded, without elongate rays; caudal fin subequal or slightly shorter than head(caudal-fin length 79.8–103.9% of head length).
Pectoral fin almost symmetrical dorsoventrally,
moderately long, typically extending to or beyond a vertical line through anal-fin origin in G.
lipi, while not extending to a vertical line throughanal-fin origin in G. arenicola. Pelvic fins fused
medially by well-developed interradial membrane; weak pelvic frenum with smooth posterior
margin present (G. arenicola) or absent (G. lipi);
pelvic fin moderately long, extending posteriorly
beyond anus.
Scales on body ctenoid with peripheral cteni;

126

K. Shibukawa and A. Iwata

Fig. 1. Ventral (top), lateral (middle) and dorsal (bottom) views of heads of 2 species of Grallenia, showing
cephalic sensory-canal pores (indicated by uppercase letters, except for AN and PN) and papillae (indicatedby roman lowercase letters). A: Grallenia arenicola, NSMT-P 73195, holotype, male, 19.4 mm SL; B: Grallenia lipi, NSMT-P 56840, holotype, male. AN and PN, anterior and posterior nares, respectively. Arrows
show positions where gill membrane attaches to isthmus. Left-side pore E of holotype of G. arenicola is abnormally undeveloped. Bars indicate 1 mm. Drawn by K. Shibukawa.

head, nape, breast, belly, anterodorsal part ofbody around base of first dorsal fin and pectoralfin base (and also, in G. lipi, trunk and anterior
half of caudal part of body, except for isolated
patch of scales just behind pectoral-fin base)naked; no modified scales with enlarged cteni.
Teeth on jaws unicuspid, slender, more or lessinwardly curved; on each jaw, 3 rows of teeth anteriorly, narrowing to a single row posteriorly;
largest teeth on anterolateral part of middle row
in lower jaws; no enlarged stout canine-like teeth
on jaws; no teeth on vomer or palatine.
Cephalic sensory systems are illustrated inFig. 1. Anterior oculoscapular canal present [G.
arenicola, with pore C (unpaired), D (unpaired),
E and F (note. —pore E of left side of the holotype abnormally absent)] or absent (G. lipi); posterior oculoscapular canal and preopercular canalabsent. Sensory papillae rows on cheek relatively
reduced, with no distinct transverse rows; all sensory-papillae rows on head uniserial or comprising a single papillae; sensory-papillae row acomprising 3 well-spaced papillae; row b short,comprising 3 papillae, its anterior end just below
anterior end of row a; row c comprising 4 papil-lae, its posterior end reaching to a vertical line
through anterior ends of rows a and b; row cp
comprising a single papilla; row d very short,comprising 2–4 papillae along jaws; row e greatly reduced (i.e., anterior end of row e locatedaround posterior corner of jaws in G. arenicola,and row e comprises only 2 papillae around posteroventral part of preopercle in G. lipi); a pair ofsensory papillae (row f ) just behind chin.
Selected osteology. Vertebrae 101718
2728; P-V typically 3/II II I I 0/9; anal-fin
pterygiophores anterior to first haemal spine 2;
single epural; posterior 3 preural vertebrae withantero-posteriorly broadened, flat neural/haemal
spines; first and second pterygiophores of second
dorsal fin lacking autogenous middle radials; ossified scapula relatively reduced, partially encircling scapular foramen; 4 pectoral radials; dorsalmost pectoral radial abuts dorsal part of cleithrum; dorsal/ventral postcleithra absent; lower
hypural plate (hypurals 12) articulates withurostyledorsal hypural plate (hypurals 34); 5
branchiostegal rays; no ossified gill rakers; basihyal triangular, its anterior margin not concave;
infrapharyngobranchial 1 absent; interarcual car-

New Goby Genus from Western Pacifictilage small, ovoid, well separated from infrapharyngobranchials; basibranchial 1 cartilaginous; basibranchial 3 reduced and very short,well separated from basibranchial 4; rostral cartilaginous; ascending process of premaxilla long,well differentiated from articular process;mesopterygoid absent; bony canal support of preopercle undeveloped; posteromedial part of preopercle with a well-developed, dorsally directedbony projection, articulating with condyle of hyomandibular for opercle.
Comparison. Grallenia clearly belongs to
the gobiid subfamily Gobiinae sensu Pezold(1993) in having: interorbital portion of oculoscapular canal fused, single unpaired anterior
interorbital pore (pore C) present (althoughcephalic sensory canals absent in Grallenia lipi);
single epural; typically 3-22110 first dorsal-fin
pterygiophore insertion pattern (3/II II I I 0/9
P-V pattern); 2 anal-fin pterygiophores anterior
to first haemal spine. The vertebral count (i.e.,
1017182728) of Grallenia places it in
the gobiine subgroup “Bathygobius Group,”
which is a phenetically-united assemblage notsupported by any synapomorphies (Birdsong et al.,1988). As well as Grallenia, the following 10
genera are assigned to this subgroup (Birdsong
et al., 1988; Hoese and Larson, 2005): Bathygobius Bleeker, 1878, Caffrogobius Smitt, 1900,
Coryogalops Smith, 1958, Glossogobius Gill,
1859, Gorogobius Miller, 1978, Hetereleotris
Bleeker, 1874, Lesueurigobius Whitley, 1950,
Nematogobius Boulenger, 1910, Psammogobius
Smith, 1935, and Palutrus Smith, 1959. [Note.—
Birdsong et al. (1988) listed the species of
Coryogalops, i.e., C. sordida, in Monishia Smith,1959 (a junior synonym of Coryogalops). The recently described genus Pascua Randall, 2005,
also assigned to the Bathygobius Group, wassubsequently regarded as a junior synonym of
Hetereleotris by Hoese and Larson (2005)].
Grallenia is readily distinguished from the othermembers of the “Bathygobius Group” in having
87 segmented caudal-fin rays and small-sized(less than 23 mm SL) slender body with a pair ofcharacteristic black spots at dorsal and ventral

127

corners of end of caudal peduncle; the othermembers of the Bathygobius Group have 98segmented caudal-fin rays and more or less larger
and stouter body, and no such black markings on
caudal-fin base. Furthermore, Bathygobius has
free filamentous rays on upper part of pectoral fin(vs. no free rays in Grallenia), small fleshy bump
below anterior naris (vs. absent), and well-developed longitudinal pattern of sensory-papillae
rows on cheek, with long parallel-running horizontal rows b and c (vs. sensory-papillae rows on
cheek relatively reduced, with short rows b and c
not running parallel one another); Caffrogobius,
Coryogalops, Gorogobius, Hetereleotris (exclusive of the Pascua-type species) and Nematogobius have well-developed, distinct transverse pattern of sensory-papillae rows on cheek (Miller,
1978, 1986; Akihito and Meguro, 1981; Hoese,
1986a, b; Harrison, 1990; Goren, 1991); the Pascua-type species of Hetereleotris [i.e., H. caudolinea (Randall, 2005), H. readerae Hoese andLarson, 2005, and H. sticta Hoese and Larson,2005] have reduced sensory-papillae rows on
cheek, but have characteristic modified scaleswith enlarged cteni on caudal-fin base (vs. absent), sensory-papilla row b closer to posterior
margin of preopercle than anterior end of row a(vs. closer to anterior end of row a than posterior
margin of preopercle), and, like all other congeners, the first gill slit closed by membrane (vs.
well open) (Randall, 2005; Hoese and Larson,2005); Glossogobius and Psammogobius havemore or less broader gill opening [extending anteriorly to, or beyond, a vertical line through posterior margin of preopercle vs. restricted (ornearly so) to pectoral-fin base], well-developed
longitudinal pattern of sensory-papillae rows on
cheek, with long parallel-running horizontal rowsb and c (exclusive of Glossogobius circumspectus with numerous transverse rows of sensory
papillae below eye) (vs. rather reduced, withshort rows b and c not running parallel one another) (Akihito and Meguro, 1975; Akihito et al.,2002); Lesueurigobius has well-developed longitudinal pattern of sensory-papillae rows on
cheek, with long parallel-running horizontal rows

128

K. Shibukawa and A. Iwatab and c (vs. rather reduced, with short rows b andc not running parallel one another), and 12–17
dorsal/anal-fin segmented rays (vs. 7–10) (Miller,1986); and Palutrus has rounded or truncate
tongue (vs. distinctly concave) (Smith, 1959;
Larson and Murdy, 2001). Other than Grallenia,
5 genera of the Bathygobius Group, i.e., Bathygobius, Glossogobius, Hetereleotris, Palutrus andPsammogobius, are distributed in the western Pacific (e.g., Larson and Murdy, 2001), but the gobies of these 5 genera are estuarine and/or reefdwelling species, not inhabiting the flat, fine andclean sandy bottoms around coral reefs as does
Grallenia.
General physiognomy of Grallenia is similar
to the Indo-Pacific commensal gobies of the genera, e.g., Bryaninops Smith, 1959, Pleurosicya
Weber, 1913, Luposicya Smith, 1959, Phyllogobius Larson, 1986, or Minysicya Larson, 2002.
Species of these genera are small (usually lessthan 40 mm SL), subtranslucent (when alive) andmore or less slender, with a pointed head, moderate to large eyes, 6 spines on first dorsal fin, typical P-V pattern of 3/II II I I 0/9, single epural, 2anal-fin pterygiophores anterior to first haemal
spine, and relatively reduced sensory-papillae
rows on cheek (Larson, 1985, 1986, 1990, 2002;
Birdsong et al., 1988). Grallenia is, however, notcommensal with any marine organisms, andreadily distinguished from these 5 genera in having: no distinct fleshy lobes developed around
pelvic-fin spine (vs. well-developed fleshy lobes
around pelvic-fin spines in the latter); 87 segmented caudal-fin rays (vs. typically 98); 10
17182728 vertebrae (vs. 101626); apair of diagonal, elongate black markings at dorsal and ventral corners of caudal-fin base (vs. nosuch markings, except for Minysicya). The lastfeature, i.e., a pair of diagonal black markings atbase of caudal fin, would be useful in distinguishing Grallenia from other similar sandy-bottom dwelling gobies (particularly young or juvenile).
Etymology. The new generic name, Grallenia, is named for Gerald R. Allen (GR
Allensuffix “-ia”), a former curator of fishes in

the WAM, in honor of his great contribution to
our knowledge of the diversity of coral-reef fishes. The gender is feminine.
Remarks. Most gobioids have 17 segmented
caudal-fin rays, exclusive of some examples with
16 or fewer segmented caudal-fin rays. Regarding
the Gobiidae, Larson (2001) noted that Nesogobius has 13 segmented caudal-fin rays, and 6 gobionelline genera [i.e., Calamiana, Chlamydogobius, Mugilogobius, Pseudogobius, Tamanka andWeberogobius (synonymized with Mugilogobius
in the latter part of her paper)] have 16 segmented caudal-fin rays. Murdy and Shibukawa (2003)
reported that Caragobius, belonging to the gobiid
subfamily Amblyopinae, has 12–15 (typically 13)
segmented caudal-fin rays. So far as we areaware, there are no gobiid genera that typicallyhave 15 segmented caudal-fin rays, exclusive of
Grallenia. Although fishes of the eleotrid subfamily Eleotrinae and the butine genus Kribia
share the condition of 15 segmented caudal-fin
rays with Grallenia (Hoese and Gill, 1993), thisis a homoplastic condition, considering that theseeleotrids lack several gobiine synapomorphies(e.g., five branchiostegal rays, pelvic fins fused
medially, single epural, and interorbital portionof oculoscapular canals unified medially) foundin Grallenia.
Within the Gobiinae, Grallenia is specialized
in having putative apomorphies such as 15 segmented caudal-fin rays, greatly reduced sensorypapillae row e, and a pair of diagonal black
markings at base of caudal fin; these characters
support the monophyly of Grallenia. However,the phylogenetic position of Grallenia is notclear as our study did not hypothesize any outgroups or sister groups. Further studies, including molecular analysis, are necessary in order toelucidate a phylogeny of Grallenia.
Grallenia arenicola sp. nov.
(New Japanese name: Sazare-haze)(Figs 1A, 2A, 3A–B and 4A; Table 1)
Gobiidae, indet. gen. and sp. 11: Senou et al., 2004: 477(underwater photograph and brief description; Kochi

New Goby Genus from Western Pacific129

Fig. 2. Schematic illustrations of 2 species of Grallenia. A: Grallenia arenicola, NSMT-P 73195, holotype,male, 19.4 mm SL; B: Grallenia lipi, one of AMS I. 43870-001, paratype, male, 16.7 mm SL.

Prefecture of Shikoku and Okinawa-jima Island of
Ryukyu Islands, Japan).
Holotype. NSMT-P 73195, male, 19.4 mm SL, off
Aka-todai Lighthouse, Kashiwa-jima Island, Kochi Prefecture, Japan, 15–20 m depth, 19 Apr. 1992 (collected by
N. Ohnishi and T. Hirata).
Paratypes. Total 12 specimens, 13.9–22.6 mm SL:
AMS I.43860-001, 4 specimens (3 males and 1 female),
13.9–15.7 mm SL, off Matsumoto-waku, Kashiwa-jimaIsland, Kochi Prefecture, Japan, 28–30m depth, 27 Nov.
1991; KPM-NI 16707, 2 specimens (females), 21.5–
22.6 mm SL, off Matsumoto-waku, Kashiwa-jima Island,
Kochi Prefecture, Japan, sandy bottom, 25 m depth, 4
May, 1993; NSMT-P 73196, 1 specimen (cleared andstained), female, 20.1 mm SL, collected with holotype;
NSMT-P 73198, 1 specimen, female, 19.7 mm SL, off
Aka-todai Lighthouse, Kashiwa-jima Island, Kochi Prefecture, Japan, 14 Aug. 1991; NSMT-P 73199, 1 specimen, female, 16.4 mm SL, off Matsumoto-waku, Kasihwajima Island, Kochi Prefecture, Japan, 12 Aug. 1992;
NSMT-P 73200, 1 specimen, female, 16.6 mm SL, Kashiwa-jima Port, Kashiwa-jima Island, Kochi Prefecture,
Japan, 10 Aug. 1992; OMNH-P 31811, 2 specimens (females), 20.3–20.8 mm SL, collected with holotype.
Non-type specimens. NSMT-P 73197, 2 specimens,
sex undetermined, 9.4–11.9 mm SL, collected with AMS
I.43860-001; NSMT-P 73201, 2 specimens (stained withalizarin red), male and female, 13.3–15.9 mm SL, off
Aka-todai Lighthouse, Kashiwa-jima Island, Kochi Prefecture, Japan, 25 Nov. 1991.

Diagnosis. Grallenia arenicola is readily
distinguished from the only other congener, G.
lipi, in having the following features: second dorsal-fin rays I, 7–8, usually I, 7 (vs. I, 9–10 in G.
lipi); anal-fin rays I, 7–8, usually I, 7 (vs. I, 9);vertebrae 10171827, usually 101727(vs. 1017182728, usually 101828);
dorsal-fin spines not elongated in either sex (vs.
first spine of first dorsal-fin greatly elongated orfilamentous in males); second dorsal fin notreaching caudal-fin base when adpressed, lengthof longest ray 14.0–17.6% of SL (vs. usuallyreaching to, or beyond, caudal-fin base when adpressed in adult, length of longest ray 17.5–
21.8% of SL); length of longest ray of anal fin
8.9–13.3% of SL (vs. 12.6–18.1% of SL); scales
on body extending anteriorly to just behind pectoral-fin base (vs. scales on body restricted on
posterior half of body, except for isolated patchof scales just behind pectoral-fin base); weak,
low pelvic frenum (vs. pelvic frenum absent);
length of pectoral fin 20.7–25.9% of SL (vs.
23.2–30.6% of SL); caudal-peduncle length
24.6–28.6% of SL (vs. 20.3–23.6% of SL);
cephalic sensory canals with pores C (unpaired),D (unpaired), E and F (vs. no cephalic sensory
canals and associated pores); no orange line on

130

K. Shibukawa and A. Iwata

Fig. 3. Preserved specimens of 4 species of Grallenia. A: Grallenia arenicola, NSMT-P 73195, holotype, male,
19.4 mm SL, Kashiwa-jima Island, Kochi Prefecture, Japan; B: Grallenia arenicola, one of OMNH-P 31811,
paratype, female, 20.8 mm SL, collected with holotype; C: Grallenia lipi, NSMT-P 56840, holotype, male,
15.5 mm SL, Ambon Island, Indonesia; D: Grallenia lipi, one of NSMT-P 56841, female, 16.3 mm SL, collected with holotype.

dorsal fins (vs. orange line at middle of dorsal
fins in life).
Description. Dorsal-fin rays VI-I, 7* (11) or
VI-I, 8 (2); anal-fin rays I, 7 (11) or I, 8* (1);pectoral-fin rays 15 (5), 16* (19) or 17 (2);pelvic-fin rays I, 5* (26); segmented caudal-fin
rays 77 (1) or 87* (12), including 55 (1),65* (10) or 75 (1) branched rays; upper unsegmented caudal-fin rays 7* (3), 8 (6) or 9 (3);lower unsegmented caudal-fin rays 5* (1), 6 (5),7 (5) or 8 (1); longitudinal scales 25 (1), 26 (3),
27 (10), 28* (9) or 29* (3); transverse scales
from anal-fin origin upward and forward to baseof first dorsal fin 4 (1), 5 (3), 6* (10) or 7 (12);transverse scales from anal-fin origin upward andbackward to base of second dorsal fin 7* (7), 8*(18) or 9 (1); transverse scales from origin of
second dorsal fin downward and backward tobase of anal fin 6 (1), 7* (14) or 8* (11); predorsal scales 0* (13); circumpeduncular scales 12*(13); gill rakers on outer surface of first arch 0
6 (1), 13* (1), 14 (2), 15 (2), 25 (3) or
27 (1); pseudobranchial filaments 4 (9) or 5*(1); vertebrae 101727* (10) or 101828(2); P-V 3/II II I I 0/9* (10) or 3/II III 0 I 0/9 (2);2* (12) anal-fin pterygiophores anterior to firstNew Goby Genus from Western Pacific131

Fig. 4. Underwater photographs of 2 species of Grallenia. A: Grallenia arenicola, Kashiwa-jima Island, Kochi
Prefecture, Japan (photographed by H. Miyamoto); B: Grallenia cf. lipi (see “Remarks” of G. lipi), Tulamben, Bali Island, Indonesia (photographed by T. Kozawa).

haemal spine; epural 1* (8).
Color when alive (based on underwater photographs, e.g., Fig. 4A). Ground color of headpale, somewhat translucent; dorsal surface ofsnout and eyes with some or several minute black
dots; iris light yellowish orange, becoming paledorsally and ventrally; ground color of body subtranslucent, with bright white belly; posterior
half of side of belly with 2 small patches ofmelanophores; a series of ca. 5 minute black dots
along lateral midline of caudal part of body; 5
short T-shaped subcutaneous black markings
along vertebral column, first one above pectoralfin base, second one at middle of trunk, third one
above anus, 4th one below end of second dorsalfin base, and fifth and posteriormost one at middle of caudal peduncle; interspace between each
black markings along vertebral column brightwhite; 5 minute black spots along midline fromanal-fin base to end of caudal peduncle; a pair ofsmall diagonally-elongate black spots at dorsal
and ventral corner of end of caudal peduncle; allclusters of melanophores on head and body overlapping similar-sized clusters of orange brown(dorsally) to pale orange (ventrally) chromatophores, except for black dots on dorsal surface of eye; ground color of fins subtranslucent;dorsal and anal fins, pectoral-fin base and middleof pelvic fin with numerous bright white speckling.
Color in alcohol. Ground color of head andbody pale yellow; minute melanophores scatteredon occipital, suborbital and postorbital areas andanterior part of operculum; eye blackish, withseveral small black dots on dorsal surface ofcornea; breast weakly (in female) or rather heavily (in male) pigmented by melanophores; several
or more dense melanophores on anteroventral
part of pectoral-fin base; a series of ca.13–14
black dots along dorsal profile of body from nape

132

K. Shibukawa and A. Iwata

to posterior part of caudal peduncle; a series ofca. 5–7 minute black dots along midlateral line oftail part of body; ca. 5–6 black dots along ventral
profile of body from anal-fin origin to posterior
part of caudal peduncle; a pair of distinct, black
diagonally-elongate black spots at dorsal andventral corners of end of caudal peduncle; dorsal
fins translucent, weakly pigmented at ventral half(melanophores usually forming a longitudinal
midline at least in second dorsal fin), with somemelanophores around tip of spine of second dorsal fin; anal fin pigmented basally and posterodistally; membrane of pelvic fin with numerous
coarse, minute melanophores, exclusive of connecting membrane between innermost rays; caudal and pectoral fins translucent. In small specimens (e.g., AMS I.43860-001), all fins are translucent, and head and body are less pigmented.
Sexual dimorphism. Male genital papillapointed, truncated in female; breast heavily pigmented with coarse melanophores in large male,weakly pigmented or pale in female and/or small
specimens.
Distribution and habitat. All examined
specimens of Grallenia arenicola were collectedfrom Kashiwa-jima Island, Kochi Prefecture,
Japan; according to Senou et al. (2004), G. arenicola (as “Gobiidae, indet. gen. and sp. 11”) is
known from Kochi Prefecture of Shikoku andOkinawa-jima Island, Ryukyu Islands, Japan.
This species is found solitary on sandy bottom
around boulders or rocky reefs at depths of 10–
50 m (Senou et al., 2004).
Etymology. The species name, arenicola, isthe combination of the Latin words “arena”(meaning “sand”) and “colo” (meaning “I inhait”),
in reference to its sandy-bottom dwelling habit.
Grallenia lipi sp. nov.
(Figs. 1B, 2B, 3C–D and 5; Table 1)
Gobiidae sp 2. Kuiter and Tonozuka, 2001: 708 (Bali, Indonesia, and Mabul, Sabah; distribution, habitat, brief
diagnosis, and 2 underwater photographs).
Holotype. NSMT-P 56840, male, 15.5 mm SL, Lilibooi, Ambon Bay, Ambon Island, Indonesia (3°45S,

Fig. 5. Freshly collected specimen of Grallenia
lipi (NSMT-P 56840, holotype, male, 15.5 mm
SL, Ambon Island, Indonesia).

128°1E), 15 m depth, 5 Dec. 1998 (collected by K. Matsuura and K. Shibukawa).
Paratypes. Total of 8 specimens, 14.6–18.8 mm SL,
collected with holotype: AMS I. 43870-001, 2 specimens,
male and female, 16.7–16.9 mm SL; KPM-NI 16708, 2
specimens, male and female, 15.5–15.8 mm SL; NSMTP56841, 3 specimens, 1 male and 2 females (1 stainedwith alizarin red), 14.6–16.4 mm SL; NSMT-P 73202, 1
specimen (cleared and stained), female, 18.8 mm SL.
Non-type specimens. NSMT-P 73203, 2 specimens,
10.8–11.2 mm SL, sex undetermined, collected with holotype.

Diagnosis. See diagnosis of congener, G.
arenicola.
Description. Dorsal-fin rays VI-I, 9 (3), VII, 10* (5) or VII-I, 10 (1); anal-fin rays I, 9* (9);pectoral-fin rays 14 (1), 15 (1), 16 (13)* or 17(3); pelvic-fin rays I, 5* (18); segmented caudalfin rays 87* (9), including 65* (9) branched
rays; dorsal unsegmented caudal-fin rays 5 (1), 6(1), 7* (5) or 8 (2); ventral unsegmented caudalfin rays 6* (7), or 7 (2); longitudinal scale rows
in anterior series 0 (1), 1 (1), 2 (2), 3* (3), 4* (2),
5 (4), 6 (2) or 7 (2); longitudinal scale rows inposterior series 10 (1), 11* (7), 12 (2), 13 (1) or
14 (5); total longitudinal scale rows 10 (1), 12(1), 13 (1), 14* (3), 15* (2), 16 (2), 17 (1), 19(4), 20 (2) or 21 (1); transverse scale rows fromanal-fin origin to base of first dorsal fin 0* (9);transverse scale rows from anal-fin origin to baseof second dorsal fin 0* (9); transverse scale rows
from second dorsal-fin origin to base of anal fin
0* (9); predorsal scales 0* (9); circumpeduncularscales 12* (9); gill rakers on outer surface of firstarch 27* (4), 28 (1), 29 (1), 38 (1) or
39 (2); pseudobranchial filamens 3 (1), 4* (7)or 6 (1); vertebrae 101727 (1) or 10
1828* (7); P-V 3/II II I I 0/9* (8) or 3/II II II I

26.3
14.6
13.3
7.0
7.6
1.5
9.5
10.2
10.9
9.5
34.1
30.3
54.7
26.3
8.1
18.9
19.4
16.0
19.2
12.8
25.9
24.0
20.9
21.6
22.3

In% of standard length
Head length
Head width
Head depth
Snout length
Eye diameter
Interorbital width
Nape width
Jaw length
Body depth
Body width
Predorsal length
Prepelvic length
Preanal length
Caudal-peduncle length
Caudal-peduncle depth
Length of D1 base
Length of D2 base
Length of longest ray of D2
Length of A base
Length of longest ray of A
P1 length
P2 length
Length of fourth segmented ray of P2
Length of fifth segmented ray of P2
C length26.3–29.8
14.6–15.2
12.6–13.3
6.7–7.7
7.6–9.0
1.0–1.5
9.5–11.3
10.2–11.5
10.9–13.3
9.5–11.0
34.0–35.0
30.1–31.7
54.0–56.1
26.2–28.6
7.6–8.1
16.4–18.9
18.4–20.4
14.0–17.4
16.5–19.2
8.9–12.8
23.0–25.9
24.0–25.2
20.9–22.4
21.6–23.1
22.3–24.3

13.9–19.4

4 males*

25.8–28.7
13.6–16.1
12.3–13.6
6.0–7.7
7.1–7.9
1.4–1.9
9.9–11.4
10.4–12.4
11.5–13.5
11.5–13.5
34.1–36.7
29.7–31.6
55.2–59.0
24.6–27.8
7.3–8.3
15.6–19.6
17.9–20.4
14.4–17.6
16.5–18.3
10.5–13.3
20.7–23.7
21.8–26.7
19.5–25.2
20.2–24.6
19.9–23.9

14.9–22.6

9 females

26.1
16.0
14.0
6.1
8.3
1.7
11.0
11.8
11.9
9.2
33.6
28.7
53.5
23.6
8.8
17.2
25.4
20.0
25.1
17.4
28.9
26.6
25.2
22.4
26.0

15.5

Male

Holotype

26.0–27.9
14.6–16.2
13.2–14.3
5.2–6.2
8.3–9.1
1.2–1.7
11.0–13.2
11.1–11.8
11.9–14.0
9.2–11.3
33.6–36.4
28.1–29.9
53.5–54.0
22.3–23.6
8.4–8.8
17.2–19.3
24.5–27.1
19.0–21.8
22.6–25.1
12.6–18.1
28.9–30.6
25.1–26.6
23.3–25.2
22.4–24.3
24.9–27.2

14.6–16.7

4 males*

27.3–29.2
15.4–17.7
14.1–15.7
5.7–6.3
8.7–9.4
1.2–1.7
12.4–13.6
11.3–12.0
12.3–13.6
11.8–13.8
35.4–36.6
29.9–32.9
55.1–56.9
20.3–22.5
7.5–8.6
14.9–18.4
24.6–26.3
17.5–20.1
22.5–24.7
13.4–15.7
23.2–28.
21.8–26.7
19.7–24.3
18.6–23.7
22.3–23.9

15.8–18.8

5 females

Holotype8 paratypes

G. lipi sp. nov.

* Including holotype. Abbreviations: A, anal fin; C, caudal fin; D1, first dorsal fin; D2, second dorsal fin; P1, pectoral fin; P2, pelvic fin.

19.4

Standard length (mm)Male

All 13 type specimens

G. arenicola sp. nov.

Proportional measurements of two species of Grallenia gen. nov.

Holotype

Table 1.

New Goby Genus from Western Pacific
133

134

K. Shibukawa and A. Iwata

0/9 (1); 2* (9) anal-fin pterygiophores anterior to
first haemal spine; 1* (9) epural.
Color when alive (based on underwater photographs, e.g., Kuiter and Tonozuka, 2001: 708).
Ground color of head and body subtranslucent;belly bright white; a series of ca. 5 short subcutaneous black lines along dorsal surface of vertebral column, as well as yellow or orange line; aseries of ca. 5 narrow horizontally-elongate rectangular white blotches on midlateral body; iris
pale; dorsal surface of eyes and snout with several minute black dots; vague black spots at dorsal
and ventral corners of end of caudal peduncle; anorange longitudinal line, sandwiched between
narrow white longitudinal bands, at middle ofdorsal fins; distal margin of dorsal fins white,
edged ventrally by orange-yellow; ventral, analand caudal fins tinged with orange-yellow.
Color when freshly collected specimen (Fig. 5).
Similar to live coloration, except as follow:
ground color of head and body whitish; blackspots at dorsal and ventral corners of end of caudal peduncle vivid and distinct.
Color in alcohol. Ground color of head andbody pale yellow; minute melanophores scatteredon occipital, suborbital and postorbital areas
and anterior part of operculum; eye blackish,with several small black dots on dorsal surface ofcornea; breast weakly (in female) or rather heavily(in male) pigmented by minute melanophores;
anteroventral part of pectoral-fin base pigmented
by various sized melanophores (a part ofmelanophores usually forming a small diagonally-elongate dark brown spot); a series of 10–12
black dots along dorsal profile of body from nape
to posterior part of caudal peduncle; a series ofca. 6–7 minute black dots on midlateral body inlarge specimens; ca. 5–8 black dots along ventral
profile of body from anal-fin origin to posterior
part of caudal peduncle; a pair of distinct, black
diagonally-elongate black spots at dorsal andventral corners of end of caudal peduncle; dorsal
fin translucent with an indistinct black line
at middle (submarginal part of second dorsal
fin sometimes weakly pigmented by minutemelanophores); membrane of anal and pelvic finsand ventral half or more of caudal fin with numerous coarse, minute melanophores (exclusiveof connecting membrane between innermost
rays very weakly pigmented); pectoral fin pale or
weakly pigmented by quite minute melanophores.
In small specimens, all fins are entirely translucent or very weakly pigmented, and head andbody less pigmented.
Sexual dimorphism. Male genital papillapointed, while truncated in female; first spine offirst dorsal fin greatly elongate and filamentous
in male, not elongate in female.
Distribution and habitat. All examined
specimens of Grallenia lipi were captured fromAmbon Bay in Ambon Island, Indonesia. They
were collected around an isolated, relatively large
coral patch on flat, clean and fine sandy bottom
at a depth of 15 m; the other fishes found in thesame habitat are the typical sandy-bottom
dwellers, e.g., the gobiid Fusigobius inframaculatus and the creediid Limnichthys nitidus; likethe congener G. arenicola, they did not display
symbiotic association with any other organisms(pers. obs., KS). According to Kuiter and Tonozuka (2001), this species (as “Filamented Pigmy
Sand-goby”) is widespread in Indonesia, andusually found on light coloured sand or rubble toabout 15 m depth; they also provided an underwater photograph taken at Mabul, Sabah.
Remarks. Based on underwater photographsfrom Indonesian waters, Kuiter and Tonozuka(2001) reported 2 unidentified goby species, assignable to Grallenia (as unidentified genus of
the Gobiidae). One of their species, named “Filamented Pygmy Sand-goby” or “Gobiidae sp 2,”
is very similar and likely conspecific with Grallenia lipi. The other Kuiter and Tonozuka’s(2001) species, named “Pygmy Sand-goby “or“Gobiidae sp 1,” appears to differ from typical
specimens of G. lipi in having heavily pigmented,
darkened coloration of head and body; it haslarger and more vivid black spots (especiallythose on caudal-fin base representing a pair oflarge longitudinally-elongate ovoid spots). A
species probably identical with Kuiter andTonozuka’s “Pygmy Sand-goby” was pho-New Goby Genus from Western Pacifictographed by T. Kozawa off Bali Island, Indoensia, and one of his images is presented in Fig. 4B.
The position of each cluster of melanophores issimilar to that of typical specimens of G. lipi,
their “Pigmy Sand-goby” is possibly a darkened
color variant of G. lipi, linked to the coloration ofsubstratum in the habitat. Kuiter and Tonozuka(2001: 708) noted that it is found, “Usually on
black sand flat,” whereas the lighter-colored “Filamented Pygmy Sand-goby” is found, “Usuallyon light coloured sand or rubble.” Further detailed comparisons based on actual specimens areneeded to clarify their taxonomy.
Etymology. The specific name, lipi, is derived from the abbreviation of the Lembaga Ilmu
Pengetahuan Indonesia (Indonesian Institute of
Science), referring that all examined specimensof this new species were captured during the
JSPS-LIPI cooperative research on marine science in Ambon Island in 1999.
Acknowledgments
We express our sincere thanks to the following
persons and institutions for specimen loans
and/or registration, providing the photographs,
and help in field collection: M. McGrouther(AMS); H. Senou (KPM); N. Ohnishi (Kyoto
Gakuen University, Japan); T. Peristiwardy (Lembaga Ilmu Pengetahuan Indonesia: LIPI); S.
Kimura (Mie University, Japan); K. Matsuura
and G. Shinohara (NSMT); S. Hosoya (Okinawa,
Japan); K. Hatooka (OMNH); W. Hiramatsu(Ehime, Japan); T. Hirata (Kochi, Japan). T.
Kozawa (Aichi, Japan) and Y. Miyamoto (Kanagawa, Japan) kindly provided the underwaterphotographs. E.O. Murdy (National Science
Foundation, U.S.A) read the manuscript and offered helpful comments. This study was financially supported partially by the Japan Society
for the Promotion of Science (JSPS) and theJapan Ministry of Education, Science, Sports andCulture Grant-in-Aid for Scientific Research on
Priority Areas (#319), Project “Symbiotic Biosphere: An Ecological Interaction Network Promoting the Coexistence of Many Species.”

135

Literature Cited
Akihito, Prince. 1984. Suborder Gobioidei. Pages
236–238 in H. Masuda, K. Amaoka, C. Araga, T.
Uyeno and T. Yoshino, eds. The Fishes of the JapaneseArchipelago. English text. Tokai University Press,
Tokyo.
Akihito, Prince and K. Meguro. 1975. Description of a
new gobiid fish, Glossogobius aureus, with notes on related species of the genus. Japanese Journal of Ichthyology, 22(3): 127–142.
Akihito, Prince and K. Meguro. 1981. A gobiid fish belonging to the genus Hetereleotris collected in Japan.
Japanese Jounal of Ichthyology, 28(3): 329–339.
Akihito, K. Sakamoto, Y. Ikeda, K. Sugiyama. 2002. Gobioidei. Pages 1139–1310 in T. Nakabo, ed. Fishes ofJapan with Pictorial Keys to the Species. Tokai University Press, Tokyo.
Allen, G., R. Steen, P. Humann and N. DeLoach. 2003.
Reef Fish Identification, Tropical Pacific. New World
Publications, Inc., Florida. 480 pp.
Birdsong, R.S., E.O. Murdy, F.L. Pezold. 1988. A study ofthe vertebral column and median fin osteology in gobioid fishes with comments on gobioid relationships.
Bulletin of Marine Science, 42: 174–214.
Goren, M. 1991. A new Coryogalops species (Pisces: Gobiidae) from South Africa. J.L.B. Smith Institute ofIchthyology, Special Publication, (52): 1–7.
Harrison, I. J. 1990. A revised synonymy of Nematogobius maindroni (Sauvage, 1880) (Teleostei, Gobiidae).
Cybium, 14(1): 27–35.
Hoese, D. F. 1986a. Family No. 240: Gobiidae. Pages
774–807 in M. M. Smith and P. C. Heemstra, eds.
Smith’s Sea Fishes. Macmillan South Africa, Johannesburg.
Hoese, D. F. 1986b. Descriptions of two new species ofHetereleotris (Pisces: Gobiidae) from the western Indian Ocean, with discussion of related species. J.L.B.
Smith Institute of Ichthyology, Special Publication,(41): 1–25.
Hoese, D. F. and A. C. Gill. 1993. Phylogenetic relationships of eleotridid fishes (Perciformes: Gobioidei). Bulletin of Marine Science, 52(1): 415–440.
Hoese, D. F. and H. K. Larson. 2005. Description of two
new species of Hetereleotris (Gobiidae) from the south
Pacific, with a revised key to species and synonymization of the genus Pascua with Hetereleotris. Zootaxa,(1096): 1–16.
Hubbs C. L. and K. F. Lagler. 1958. Fishes of the Great
Lakes Region. Cranbrook Institute of Science, Bloomfield Hills, Michigan. vii213 pp., 44 pls.
Kuiter, R. H., T. Tonozuka. 2001. Indonesian Reef Fishes.
Zoonetics, Australia. 893 pp.
Larson, H. K. 1985. A revision of the gobiid genus

136

K. Shibukawa and A. Iwata

Bryaninops (Pisces), with a description of six new
species. The Beagle, Occasional Papers of the Northern Territory Museum of Arts and Sciences, 2(1): 57–
93.
Larson, H. K. 1986. Phyllogobius, a new generic name for
Cottogobius platycephalops Smith (Pisces: Gobiidae),and a redescription of the species. The Beagle, Occasional Papers of the Northern Territory Museum of Arts
and Sciences, 3(1): 131–136.
Larson, H. K. 1990. A revision of the commensal gobiid
genera Pleurosicya and Luposicya (Gobiidae), with descriptions of eight new species of Pleurosicya and discussion of related genera. The Beagle, Records of theNorthern Territory Museum of Arts and Sciences, 7(1):1–53.
Larson, H. K. 2001. A revision of the gobiid fish genus
Mugilogobius (Teleostei: Gobioidei), and its systematic
placement. Records of the Western Australian Museum,
Supplement, (62): 1–233.
Larson, H. K. 2002. A new genus of small coral-reef goby
(Teleostei: Gobiidae) from the Indo-west Pacific, withdiscussion of its relationships. The Beagle, Records ofthe Museums and Art Galleries of Northern Territory,
18: 63–71.
Larson, H. K. and E. O. Murdy. 2001. Gobiidae. Pages
3578–3603 in K. E. Carpenter and V. H. Niem, eds.
FAO Species Identification Guide for Fisheries Purpose. The Living Marine Resources of the Western
Central Pacific. Volume 6. Bony Fishes Part 4 (Labridae to Latimeriidae), Estuarine Crocodiles, Sea Turtles,
Sea Snakes and Marine Mammals. FAO, Rome.
Leviton, A. E., R. H. Gibbs, Jr., E. Heal and C. E. Dawson. 1985. Standards in herpetrogy and ichthyology:
Part I. Standard symbolic codes for institutional resource collections in herpetology and ichthyology.

Copeia, 1985(3): 802–832.
Miller, P. J. 1978. The status of the west African fish Gobius nigricinctus with reference to New World autochthones and an Old World colour-analogue. Zoological journal of the Linnean Society, 64(1): 27–39.
Miller, P. J. 1986. Gobiidae. Pages 1019–1085 in P.J.P.
Whitehead, M.-L. Bauchot, J.-C. Hureau, J. Nielsen &E. Tortonese, eds. Fishes of the North-eastern Atlantic
and the Mediterranean. UNESCO, Paris.
Murdy, E. O. and K. Shibukawa. 2003. A revision of the
Indo-Pacific genus Caragobius (Gobiidae: Amblyopinae). Zootaxa, (301): 1–12.
Pezold, F. 1993. Evidence for a monophyletic Gobiinae.
Copeia, 1993(3): 634–643.
Potthoff, T. 1984. Clearing and staining techniques. Pages
35–37 in H. G. Moser, W. J. Richards, D. M. Cohen,
M. P. Fahay, A. W. Kendall Jr. and S. L. Richardson,
eds. Ontogeny and Systematics of Fishes. American
Society of Ichthyologists and Herpetologists Special
Publication 1.
Randall, J. E. 2005. Reef and Shore Fishes of the South
Pacific: New Caledonia to Tahiti and the Pitcairn Islands. University of Hwai’i Press. x707 pp.
Senou, H., T. Suzuki, K. Shibukawa and K. Yano. 2004. A
Photographic Guide to the Gobioid Fishes of Japan.
Heibonsha, Tokyo. 536 pp.
Smith, J. L. B. 1959. Gobioid fishes of the families Gobiidae, Periophthalmidae, Trypauchenidae, Taenioididae
and Kraemeriidae of the western Indian Ocean. Ichthyological Bulletin, (13): 185–25, pls 9–13.
Manuscript received 26 March 2006; revised 29 January2007; accepted 7 February 2007.
Associate editor: K. Matsuura.