Review of the Cheek-spine Goby Genus Gladiogobius (Actinopterygii, Perciformes, Gobiidae), with Descriptions of Two New Species from the Indo-West Pacific

2007·0.66 MB·English

by  Shibukawa, K

Bull. Natl. Mus. Nat. Sci., Ser. A, 33(4), pp. 193–206, December 21, 2007
Review of the cheek-spine goby genus Gladiogobius (Actinopterygii,
Perciformes, Gobiidae), with descriptions of two new species
from the Indo-West Pacific
Koichi Shibukawa1and Gerald R. Allen2
1Nagao Natural Environment Foundation, 3–10–10 Shitaya, Taito-ku, Tokyo 110–0004, Japan
E-mail: [email protected]
2Department of Aquatic Zoology, Western Australian Museum, Francis Street, Perth, Western Australia
E-mail: [email protected]
Abstract The Indo-West Pacific cheek-spine goby genus Gladiogobius Herre is reviewed. The
genus contains three species, G. brevispinis sp. nov., G. ensifer Herre, 1933 and G. rex sp. nov.,
which are distinguished from other gobiine genera in having the following combination of charac-
ters: a single long, posteriorly-directed, spur-like spinous projection at rear margin of preopercle;
low cutaneous ridge along predorsal midline; no scales on head and nape; distinct transverse pat-
tern of sensory-papilla rows on cheek. Gladiogobius brevispinis is described on the basis of 11
specimens (12.5–42.6mmSL) from Japan (Ryukyu Islands), China (Hainan Island) and Indonesia
(Sulawesi and Bali). It differs from congeners in having a short preopercular spine (clearly shorter
than length of preopercular canal) and no filamentous dorsal-fin spines. Gladiogobius rex is de-
scribed on the basis of 20 specimens (17.5–38.9mmSL) from Phuket, Thailand, but also occurs
widely in the Indian Ocean and the Red Sea. It is readily distinguished from congeners in having a
very long preopercular spine (extending well beyond opercular margin). A redescription is also
provided for G. ensiferbased on 17 specimens (24.3–34.0mmSL) from scattered localities in the
West Pacific. We also include a key to the species of Gladiogobius.
Key words: Gobiidae, Gladiogobius brevispinissp. nov., Gladiogobius rexsp. nov., Gladiogob-
ius ensifer, Indo-West Pacific.
Three Indo-west Pacific genera within the gob- best, due to a greatly diminished fish fauna com-
iid subfamily Gobiinae (sensuPezold, 1993) pos- pared to that of coral reefs and silty conditions
sess one or more posteriorly-directed, preopercu- that cause poor underwater visibility. However,
lar or cheek spines: AsterropteryxRüppell, 1830; on this occasion there was good tidal flushing
Gladiogobius Herre, 1933; Oplopomus Valenci- with an influx of clear water. Consequently, a
ennes inCuvier and Valenciennes, 1837. Gladio- number of interesting fishes were observed and
gobius was described by Herre (1933), based on several were captured using a small multiprong
his new species, Gladiogobius ensifer, from spear. The collection included two specimens of
Waigeo Island, western New Guinea (Raja an unfamiliar goby subsequently identified as an
Ampat Islands, Irian Jaya Barat Province, In- undescribed species of Gladiogobius.
donesia) and Culion Island in the Calamianes After the discovery of the Bali fish, GRA
Group, Philippines. The genus has been consid- learned that the first author was in the process of
ered monotypic since its original description. revising Gladiogobius, based on research involv-
During February 1999 the second author ing collections housed in several institutions in
(GRA) had a rare opportunity to dive in the inner Japan. We therefore decided to collaborate on a
estuary at Gilimanuk, off the western end of Bali, review of this genus, which now contains three
Indonesia. Estuary diving is usually mediocre at species, including two new taxa described herein.
194 Koichi Shibukawa and Gerald R. Allen
Materials and Methods count was the number of scales on the ventral
Institutional abbreviations follow Leviton et midline of the preventral area (scales are typical-
al. (1985), except for NCIP (Pusat Penelitian dan ly embedded); gill rakers, including all rudi-
Pengembangan Oseanologi, Jakarta, Indonesia). ments, were counted on the outer side of first
Species accounts are presented in alphabetical arch; counts of the pseudobranchial filaments in-
order. cluded all rudiments. Scales (except for predorsal
The methods for measurements followed those and circumpeduncular scales) and paired-fin rays
of Hubbs and Lagler (1958), with exceptions were counted on both sides, but gill rakers and
given below (the snout tip refers to the mid-ante- pseudobranchial filaments were counted on the
riormost point of the upper lip): interorbital right side only. Meristic counts of the holotype
width was the least width between the innermost are indicated with an asterisk in the species de-
rims of the right and left eyes; jaw length was scriptions. The number of paratypes having a
measured between the snout tip and the posteri- particular count appears in parentheses. Osteo-
ormost point of the lips; length of cheek spine logical features were observed from radiographs
was the least distance between the anterior bony and cleared and stained specimens, following the
ridge of the preopercular canal support (clearly method of Potthoff (1984). The methods of Aki-
visible from external view) and tip of the spine; hito et al. (1984) were used in describing the pat-
width of nape was measured between the upper- tern of the interdigitation of the dorsal-fin ptery-
most ends of the gill openings; body depth was giophores between the neural spines (“P–V”).
measured at the anal-fin origin; preanal and pre- Cephalic sensory canals and papillae were ob-
pelvic lengths were measured from the snout tip served on specimens stained with cyanine blue,
to the origin of each fin; pectoral-fin length was and abbreviations for these followed Akihito et
measured from the base to the tip of the longest al. (1984) and Miller (1986), respectively.
ray; pelvic-fin length was measured between the
base of the pelvic-fin spine and the distal tip of
Genus GladiogobiusHerre, 1933
the longest (4th) segmented ray; height of pelvic-
[Japanese name: Togenaga-haze zoku]
fin connecting membrane was measured at the
GladiogobiusHerre, 1933:23 (type species: Gladiogobius
lowest point of the membrane (usually measured
ensiferHerre, 1933, by monotypy).
from mid-point between the right and left sides
of the innermost ray to the mid-posterior edge of Diagnosis. Gladiogobius is distinguished
the connecting membrane); caudal-fin length was from all other gobiid genera in having the follow-
measured from the base to the tip of the middle ing combination of characters: a single distinct,
caudal-fin ray (not including elongate filamen- posteriorly-directed spur-like bony spinous pro-
tous part). Measurements were made with jection at rear margin of preopercle; low cuta-
calipers under a dissecting microscope to the neous ridge along predorsal midline; no scales on
nearest 0.01mm. The methods of counts fol- head and nape; distinct transverse pattern of sen-
lowed Akihito et al. (1984), except for the fol- sory-papilla rows on cheek.
lowing: longitudinal scale count was the number Description. Dorsal-fin rays VI-I, 9–10 (usu-
of oblique rows starting from just above the dor- ally VI-I, 10); anal-fin rays I, 8–9 (usually I,
salmost attachment of the opercular membrane 9–10); pectoral-fin rays 16–18; pelvic-fin rays I,
and proceeding posteriorly to the mid-base of the 5; segmented caudal-fin rays 9(cid:1)8, including
caudal fin; three types of transverse scale count 6–7(cid:1)6–7 branched rays; dorsal unsegmented
were taken (see descriptive accounts); circumpe- caudal-fin rays 5–8; ventral unsegmented caudal-
duncular scale count was the number of zigzag fin rays 3–8 (usually 5 or more); longitudinal
rows along a vertical line around the narrowest scales 23–26; transverse scales from anal-fin ori-
point of the caudal peduncle; preventral scale gin upward and forward to base of first dorsal fin
Review of Gladiogobuius 195
9–13; transverse scales from anal-fin origin up- than second dorsal fin; first dorsal fin close to,
ward and backward to base of second dorsal fin but not connected to, second dorsal fin by mem-
8–12; transverse scales from second dorsal-fin brane; fourth (or, occasionally, fifth) spine of first
origin downward and backward to base of anal dorsal fin longest (127.4% of preceding spine in
fin 7–11; predorsal scales 0; prepelvic scales length), elongate, filamentous (if fifth spine
0–5; circumpeduncular scales 12; gill rakers longest, the fourth spine also long and filamen-
3–5(cid:1)8–10; pseudobranchial filaments 4–8. tous and only slightly shorter than fifth spine) ex-
Body moderately elongate and compressed. cept in G. brevispinis; all dorsal-fin spines slen-
Head compressed, its width 72.2–95.3% of der and flexible; all segmented rays of second
depth. Snout short, its length subequal or usually dorsal fin branched; eighth or ninth segmented
slightly shorter than eye diameter (snout length ray of second dorsal fin longest, its distal end not
71.7–113.4% of eye diameter); snout does not extending to, or beyond, a vertical through base
protrude beyond upper lip. Eye dorsolateral, of caudal fin when adpressed. Origin of anal fin
large, its diameter 22.1–31.1% of head length; on a vertical with base of second segmented ray
interorbital space narrow, its width narrower than of second dorsal fin; height of anal fin slightly
pupil diameter and 2.4–6.0% of head length. Dis- lower than second dorsal fin; anal-fin spine slen-
tinct, low cutaneous ridge along predorsal mid- der and flexible; all segmented anal-fin rays
line, extending forward beyond a vertical through branched; eighth (in G. ensifer and G. rex) or
preopercular margin (but not reaching to eyes). ninth (in G. brevispinis) segmented ray of anal fin
Gape oblique, forming an angle of about 25–40 longest, its distal end not extending to, or slightly
degrees with body axis. Lower jaw subequal to beyond, a vertical through base of caudal fin
upper jaw; posterior end of jaws reach to below when adpressed. Caudal fin rounded or nearly
pupil; jaw length 29.0–36.8% of head length. truncate; tenth (and, occasionally, eighth and
Anterior nasal opening a short tube, slightly pro- ninth) segmented ray slightly elongate and fila-
truding over upper lip; no fleshy flap at tip of an- mentous. Pectoral fin nearly lanceolate, reaching
terior naris; posterior nasal opening a simple posteriorly to a vertical line through base of first,
pore with only a slightly elevated anterior mar- second or third segmented ray of second dorsal
gin, located closer to eye than anterior naris. Tip fin; all pectoral-fin rays branched, excluding sim-
of tongue nearly truncate, free from floor of ple uppermost 1–2 and, sometimes, ventralmost
mouth. Posteroventral margin of lower lip inter- ray Origin of pelvic fin below or only a slightly
rupted at symphysis. Mental flap on chin unde- anterior to a vertical line through origin of first
veloped. Gill opening moderately narrow, not ex- dorsal fin; pelvic fins almost separated, fused me-
tending anteriorly to a vertical line through pre- dially via very low connecting membrane (its
opercular margin; gill membranes attach to isth- height 11.2–28.6% length of fifth segmented
mus. No fleshy praojections on lateral wing of pelvic-fin ray) between innermost rays; no
shoulder girdle. A distinct, posteriorly-directed, pelvic-fin frenum; all segmented pelvic-fin rays
spur-like spinous bony projection at posterior branched; fourth segmented ray longest; fifth
margin of preopercle. Gill rakers on outer surface segmented ray 61.6–87.7% of preceding (fourth)
of ventral arm of first arch well developed, long ray in length; pelvic fin reaching posteriorly to
and thin (i.e. blade-like); rakers on outer surface base of spine or first or second segmented rays of
of dorsal arm of first arch well developed, but anal fin when adpressed.
distinctly shorter than neighbor rakers on ventral Most of body and caudal-fin base covered by
arm; first gill slit well developed. Caudal pedun- ctenoid scales with peripheral cteni, except belly,
cle moderately slender, its depth 50.7–67.4% of pectoral-fin base, and prepelvic region with cy-
caudal-peduncle length. First dorsal fin (exclud- cloid scales (pectoral-fin base naked in one
ing filamentous parts) subequal or slightly higher species); head and nape naked.
196 Koichi Shibukawa and Gerald R. Allen
Fig. 1. Heads of three species of Gladiogobius, showing cephalic sensory-canal pores (indicated by uppercase
letters, except for AN and PN) and sensory papillae (indicated by lowercase letters). A) Gladiogobius ensifer,
NSMT-P 61522, male, 24.8mmSL; B) Gladiogobius brevispinis, NCIP 6186, holotype, male, 42.6mmSL;
C) Gladiogobius rex, NSMT-P 44892, paratype, male, 31.6mmSL. AN and PN, anterior and posterior nares,
respectively. Arrows show the position where the gill membrane is attached to the isthmus. Bars indicate 3
mm. Drawn by K. Shibukawa.
Teeth in both jaws slender, unicuspid; each jaw vertical row; row brelatively short, not extending
with 3–5 rows of teeth anteriorly, narrowing to below eye; rows e and i broadly interrupted in
single row posteriorly; teeth on outermost row middle section; a pair of short longitudinal rows
slightly enlarged, larger than those of inner rows; of sensory papillae just behind chin ((cid:3)row f);
no stout canine-like teeth on jaws; no teeth on row n a short transverse row; row s2 a short lon-
vomer or palatine. gitudinal row comprising 1–3 papillae. Sensory
Cephalic sensory systems are illustrated in papillae on midlateral body forming uniserial
Fig. 1. Anterior oculoscapular-canal pores B(cid:2), C vertical rows, each restricted to a single scale;
(unpaired), D (unpaired), E, F, G and H(cid:2); posteri- three radiating rows of sensory papillae on cau-
or oculoscapular-canal pores K(cid:2)and L(cid:2); preoper- dal fin along the sixth, ninth and twelfth (or
cular-canal pores M(cid:2), N and O(cid:2); right and left eleventh) segmented caudal-fin rays.
sides of anterior oculoscapular canals fused me- Selected osteology. Vertebrae 10(cid:1)16(cid:3)26;
dially in interorbital space. All cephalic sensory- each epineural not fused to relevant pleural; P-V
papillae rows uniserial or comprising a single 3/II II I I 0/9; anal-fin pterygiophores anterior to
papilla, not forming multiple lines or aggrega- first haemal spine 2; single epural; first and sec-
tions; distinct transverse pattern of sensory papil- ond pterygiophores of second dorsal fin lacking
lae rows on cheek; six short transverse rows of autogenous middle radials; ossified scapula mod-
sensory papillae (rows 1–6) below eye; rows 4 erately developed, encircling scapular foramen
and 5 sometimes continuous, forming a single dorsally; 4 pectoral radials; dorsalmost pectoral
Review of Gladiogobuius 197
radial abuts dorsal part of cleithrum; short rib- rigid and pungent first spine of first and second
like ventral postcleithra present; lower hypural dorsal fins (vs. flexible), and entirely united
plate (hypurals 1(cid:1)2) articulates with urostyle (cid:1) pelvic fins (vs. largely separated).
dorsal hypural plate (hypurals 3(cid:1)4); dorsal and The species of Gladiogobiusare small bottom-
ventral procurrent cartilages moderately short, dwellers (largest specimen examined, 42.6mm
extending anteriorly to a little beyond distal tips SL), typically found in shallow sheltered bays
of neural and haemal spines of 14th caudal verte- and lagoons with sandy-mud bottoms.
bra, respectively; 5 branchiostegal rays; gill rak- Included species. Gladiogobius comprises
ers on outer side of first arch slender, short and three species: G. brevispinis sp. nov., G. ensifer,
filamentous; other gill rakers short and stout, and G. rexsp. nov.
each bearing short distal spines; no gill rakers on
inner and outer sides of first and second arches,
Key to the species of Gladiogobius
respectively; basihyal triangular, its anterior mar-
gin not concave; infrapharyngobranchial 1 ab- 1a. Cheek spine very long, extending posterior-
sent; interarcual cartilage relatively long corn- ly to well beyond posterior margin of oper-
like; basibranchial 1 cartilaginous; basibranchial cular membrane (Fig. 1C), its length about
3 moderately long, approaching basibranchial 4 twice eye diameter; several distinct, minute
posteriorly; rostral cartilaginous; ascending black (dark brown when alive) dots across
process of premaxilla long, well differentiated cheek from posterior end of jaws to base of
from articular process; mesopterygoid absent; cheek spine (Red Sea, Indian Ocean, An-
posteroventral portion of preopercle with a flat- daman Sea and northwestern Java, Indone-
tened, stout, blade-like, posteriorly-directed spin- sia) ........................................G. rexsp. nov.
ous projection; bony canal support developed 1b. Cheek spine not extending posteriorly to
throughout vertical portion of preopercle; anteri- posterior margin of opercular membrane
or margin of middle of preopercle without a dor- (Figs. 1A & 1B), its length subequal or less
sally directed bony projection; anterior margin of than eye diameter; no distinct minute black
vomer rounded; posterior laminar portion of dots across cheek (usually single or some
mesethmoid well ossified, forming interorbital faint, small dusky blotches present just be-
septum anteriorly; frontals not fused medially; a hind posterior end of jaws). ......................2
low mid-dorsal bony crest on posterior part of 2a. Cheek spine relatively long, its length
supraoccipital. 21.3–26.7% of head length and usually
Sexual dimorphism. Male genital papilla nar- subequal to distance between preopercular-
row and pointed, broad and rounded in female. canal pores M(cid:2) and O(cid:2) (Fig. 1A); typically
Remarks. As noted above, three Indo-Pacific nine segmented anal-fin rays; fourth spine
gobiine genera possess one or more posteriorly- of first dorsal fin greatly elongate, filamen-
directed preopercular spines, i.e., Asterropteryx, tous, longer than preceding spine; base of
Gladiogobius, and Oplopomus. Gladiogobius is pectoral fin broadly covered by embedded
readily distinguished from the other two genera cycloid scales; dusky marking just above
in having a low cutaneous ridge along the predor- dorsal end of gill opening ovoid, distinctly
sal midline and a pair of conspicuous black spots deeper than preceding dusky marking
just behind the isthmus. In addition, Aster- (Ryukyu Islands of Japan, Palau, Solomon
ropteryx differs in having scales on cheek, oper- Islands, Philippines, Indonesia, Gulf of
culum and nape (vs. head and nape naked in Thailand, and northern Australia)................
Gladiogobius), and Oplopomus has multiple lon- .....................................................G. ensifer
gitudinal pattern of sensory-papillae rows on 2b. Cheek spine short, its length 12.4–17.9% of
cheek (vs. transverse pattern in Gladiogobius), head length and distinctly shorter than dis-
198 Koichi Shibukawa and Gerald R. Allen
tance between preopercular-canal pores M(cid:2) Nobaru-zaki Point, Iriomote-jima Island, Ryukyu
and O(cid:2) (Fig. 1B); typically 10 segmented Islands, Japan, 14 Aug. 1996 (collected by H.
anal-fin rays; no elongated and filamentous Yoshigou); WAM-P.31531.004, 1 specimen (fe-
spines on first dorsal fin (some middle male), 41.9mmSL, collected with holotype.
spines may be weakly filamentous, but Non-type material. NSMT-P 76626, 1 speci-
fourth spine always subequal in length to men (juvenile), 14.5mmSL, Sendakan, west
preceding spine); no scales on pectoral-fin coast of Lembeh Island, off Sulawesi, Indonesia,
base, except some embedded, small cycloid 1m depth, 17 July 2000 (collected by K.
scales on at ventral part; dusky marking Shibukawa).
just above dorsal end of gill opening usual- Diagnosis. Gladiogobius brevispinis is dis-
ly narrow, dash-like, subequal in depth to tinguished from other congeners in having: a
preceding dusky marking (Ryukyu Islands short cheek spine (its length 3.7–5.7% and
of Japan, Hainan Island, Yap, and Indone- 12.4–17.9% in SL and head length, respectively,
sia)............................G. brevispinissp. nov. vs. 6.5–16.4% and 21.3–53.1% in SL and head
length, respectively, in other congeners); typical-
ly 10 segmented anal-fin rays (vs. typically 9); no
Gladiogobius brevispinissp. nov.
elongate, filamentous spines on first dorsal fin
(Figs. 1B, 2A, 3A) (except tip of middle spines occasionally fila-
[New Japanese name: Kotoge-haze]
mentous, but fourth spine subequal in length to
[New English name: Short-spined Goby]
preceding spine vs. at least fourth spine well
Gladiogobius ensifer (non Herre); Masuda and elongate, filamentous, longer than preceding
Kobayashi, 1994: 329, fig. 1 (Yaeyama Islands); Akihi- spine); no scales on pectoral-fin base, except
to et al., 1993: 1012 (line drawing of whole body); Ak-
some embedded, small cycloid scales on ventral
ihito et al., 2000: 1238 (line drawing of whole body);
part (vs. largely covered by embedded cycloid
Akihito et al., 2002: 1238 (line drawing of whole
body); Kuiter and Tonozuka, 2001: 676, fig. A (Java, scales); dusky marking just above dorsal gill at-
Indonesia). tachment usually narrow, dash-like, subequal in
Gladiogobius sp. 1; Senou et al., 2004: 391 (Iriomote- depth to preceding dusky marking (vs. ovoid, dis-
jima Island, Yaeyama Islands of Ryukyu Archipelago,
tinctly deeper than preceding dusky blotch).
Japan).
Description. The following description is
Holotype. NICP 6186, male, 42.6mmSL, based on the type series. Dorsal-fin rays VI-I,
Gilimanuk, Bali, Indonesia (08°12(cid:2)S, 114°27(cid:2)E), 10* (9); anal-fin rays I, 9 (1) or I, 10* (8); pec-
inner lagoon next to mangrove, 1.0–1.5m depth, toral-fin rays 16* (6) or 17* (12); pelvic-fin rays
7 Feb. 1999 (coll. by G. R. Allen). I, 5* (18); segmented caudal-fin rays 9(cid:1)8* (7),
Paratypes. Nine specimens, 12.5–41.9mm including 6(cid:1)6 (1), 6(cid:1)7* (4) or 7(cid:1)7 (2)
SL : NSMT-P 55899, 2 specimens (2 males, in- branched rays; dorsal unsegmented caudal-fin
cluding 1 cleared and stained), 26.7–28.1mmSL, rays 5 (1), 6 (1) or 7* (3); ventral unsegmented
Luhuitou Bay, Hainan Island, China, 3 Dec. 1997 caudal-fin rays 5 (2), 6* (1) or 7 (2); longitudinal
(collected by G. Shinohara); URM-P 4123, 2 scales 24 (4), 25* (6) or 26 (4); transverse scales
specimens (male and female), 24.1–26.9mmSL, from anal-fin origin upward and forward to base
Yonada-gawa River, Iriomote-jima Island, of first dorsal fin 12* (7) or 13 (7); transverse
Ryukyu Islands, Japan, 16 Aug. 1982 (collected scales from anal-fin origin upward and backward
by Yamamura and Totani); URM-P 31223, 1 to base of second dorsal fin 9* (3) or 10* (11);
specimen (female), 29.3mmSL, Miyako-jima Is- transverse scales from second dorsal-fin origin
land, Ryukyu Islands, Japan, 11–12 June 1993 downward and backward to base of anal fin 8*
(collected by Kanashiro); URM-P 36312, 2 spec- (1), 9* (10) or 11 (2); predorsal scales 0* (7);
imens (female and juvenile), 12.5–24.6mmSL, prepelvic scales 0* (2), 1 (2) or 2 (3); circumpe-
Review of Gladiogobuius 199
s
8 paratypes aFemale 4 specimen 17.5–38.9 29.5–32.06.4–7.96.5–8.61.0–1.69.0–10.016.6–20.014.9–18.112.0–13.816.6–20.711.3–14.334.1–36.832.2–36.157.8–63.220.6–23.912.2–13.921.2–25.123.2–25.217.4–20.127.2–32.126.7–32.125.7–30.120.6–24.424.5–29.213.2–16.4
v. 1 1
o (cid:1)
ogobius rexsp. n Holotype Males 5 specimens 30.8–38.4 29.1–30.66.5–7.36.7–7.01.0–1.58.9–9.817.8–18.814.8–16.910.5–13.019.2–20.211.9–13.633.9–34.831.6–35.258.5–62.221.9–23.412.3–13.523.5–25.123.8–25.417.5–19.927.8–34.527.9–30.927.5–28.617.3–23.824.4–28.513.5–15.9
di
a
Gl 0
5
Holotype Female MBC 166 33.1 30.9 6.7 7.6 1.3 10.0 19.5 14.9 12.7 20.7 12.8 34.1 32.8 61.7 20.7 13.0 25.1 24.9 19.3 31.2 32.1 29.6 23.9 27.2 15.1/15.0
s P
u
bi
o
ments of three species of Gladiog Gladiogobius ensifer Non-type specimens MalesFemales 5 specimens6 specimens 24.8–31.124.3–34.0 30.2–32.130.0–31.76.9–8.26.8–7.67.1–8.17.2–8.21.0–1.41.0–1.59.5–10.69.6–10.117.8–20.118.3–20.014.1–17.014.8–16.311.6–13.512.0–12.918.5–20.818.6–20.211.1–13.510.8–14.033.1–34.333.9–35.733.0–35.733.6–35.558.0–59.960.5–63.123.0–24.922.6–23.312.9–13.912.4–13.822.7–24.122.5–24.723.7–24.822.1–25.518.2–20.418.0–19.029.3–34.129.3–30.528.8–34.128.9–31.227.2–32.827.7–29.820.2–26.221.3–23.425.3–31.224.3–27.66.8–8.66.5–7.5 ue representing the left one.n; P, pectoral fin; P, pelvic fin.12
Table1.Proportional measure Gladiogobius brevispinissp. nov. (cid:1)HolotypeHolotype7 paratypes aFemalesMaleMales NCIP 61864 specimens4 specimens 42.6 26.7–42.624.1–41.9 30.7 30.7–33.229.4–33.07.0 6.6–7.06.2–7.26.8 6.8–8.07.2–8.41.8 1.6–1.80.8–1.711.3 11.2–11.310.0–11.418.9 18.9–19.518.4–20.115.6 15.6–16.014.3–16.212.5 11.9–12.511.4–13.122.1 19.1–22.119.4–21.114.5 12.5–14.512.2–15.132.0 32.0–34.134.4–36.136.9 36.9–38.035.5–40.560.0 60.0–60.259.8–63.222.4 22.0–22.420.9–22.315.1 12.1–15.112.4–13.824.0 20.9–24.020.2–23.825.0 22.8–25.021.5–23.722.0 20.4–22.019.1–20.529.8 29.8–30.728.0–32.328.9 28.9–29.826.5–29.727.9 27.5–27.925.2–28.123.9 22.8–23.921.2–22.828.2 27.7–28.225.8–28.35.2/5.54.2–5.73.7–5.5 sides of cheek spine in holotype by a slash, the first valn; C, caudal fin; D, first dorsal fin; D, second dorsal fi12
Standard length (mm) In % of standard lengthHead lengthSnout lengthEye diameterInterorbital widthJaw lengthHead depthHead widthNape widthBody depthBody widthPredorsal lengthPrepelvic lengthPreanal lengthCaudal–peduncle lengthCaudal-peduncle depthLength of Dbase1baseLength of D2Length of A basePlength1Plength2Length of 4th Pray2Length of 5th Pray2C lengthbLength of cheek spine aIncluding holotype.bMeasured bilaterally.cValues of left and right Abbreviations: A, anal fi
200 Koichi Shibukawa and Gerald R. Allen
Fig. 2. Three species of Gladiogobius. A) G. brevispinis, WAM-P.31531.004, paratype, female, 41.9mmSL; B)
G. ensifer, NSMT-P 61522, male, 24.8mmSL; C) G. rex, PMBC 16650, holotype, female, 33.1mmSL. Pho-
tographed by K. Shibukawa.
duncular scales 12* (7); gill rakers 3(cid:1)9 (1), 4(cid:1)9 short, parallel horizontal dark brownish gray
(1), 4(cid:1)10 (3) or 5(cid:1)9 (1); pseudobranchial fila- stripes from posteroventral margin of eye to a
ments 6 (1), 7 (3) or 8 (2). vertical line through posterior margin of pre-
Color when alive. The following description opecle; a single, slightly oblique, dark brownish
is based on underwater photographs taken by K. gray irregular stripe (tinged with orange) from
Yano from Iriomote-jima Island, Ryukyu Archi- posterior corner of jaws to posterior margin of
pelago, Japan (Senou et al., 2004: 391): ground operclum; ground color of eyes light gray dorsal-
color of head light bluish gray dorsally, pale ly, dark gray ventrally; short blackish transverse
beige ventrally; several minute pale sky spots lines or mottles on dorsal surface of eye; pupil
scattered on cheek and operculum, as well as a encircled by reddish yellow; two short, parallel,
short oblique, pale blue line below eye; two dash-like oblique black lines just above opercu-
Review of Gladiogobuius 201
Fig. 3. Underwater photographs of two species of Gladiogobius. A) Gladiogobius brevispinis, Yap Island, Mi-
cronesia (photographed by G. R. Allen); B) Gladiogobius ensifer, Tatsugo, Amami-oshima Island, Ryukyu Is-
lands, Japan, 2m depth (photographed by H. Kanehara). Note.—In the photo specimen of G. brevispinis(A),
fourth spine of first dorsal fin is elongate and filamentous, but subequal in length to preceding spine.
lum and pectoral-fin base; occipital region and records are expected to result from further estu-
nape with numerous dense, small irregular- arine collections, especially in areas adjacent to
shaped, dark brownish gray spots; ground color the Indo-Australian Archipelago. GRA encoun-
of body light bluish gray, except for pale beige tered about 15 individuals (including holotype
belly; a series of five horizontally elongate, dusky and a single paratype) at Bali, Indonesia in
blotches on mid-lateral body, also a pair of simi- depths of 1.0–1.5m near the mangrove fringe of
lar colored blotches on side of belly; 5–6 hori- a large enclosed estuary with the bottom consist-
zontal series of minute brownish spots on side of ing of fine sand and silt. Solitary fish or pairs
body; numerous minute pale blue dots scattered were present at the mouth of burrows that were
on body (those on belly especially conspicuous excavated under small, rocky outcrops. He also
and enlarged); fins nearly transparent, with nu- observed several individuals associated with an
merous enlarged white chromatophores (pectoral unidentifed alpheid shrimp that shared the same
and pelvic fins slightly tinged with yellow); mid- burrow. Moreover, GRA recently photographed
dle of caudal fin with ca. 4–5 indistinct vertical the species well to the east of the known distribu-
or arched reddish gray lines. Coloration of live tion at Yap Island in Micronesia, where it was
specimen from Java, Indonesia (Kuiter and living in coralline areas adjacent to coastal man-
Tonozuka, 2001: 676, photo A of “Gladiogobius groves (Fig. 3A). Senou et al. (2004) reported
ensifer”), is similar to the Japanese specimens this species from the Yaeyama Group, Ryukyu
described above, except the ground ground color Islands (Ishiagki-jima Island and Iriomote-jima
of the head and body is paler. Island) where it was found solitary in estuaries
Color in alcohol. Similar to live coloration and inner reef areas with sand-mud bottoms and
except the blue spots and lines on the head and dead-coral rubble in less than 2 m depth.
body are faded as well as the white and/or yellow Remarks. Akihito et al. (1993, 2000, 2002)
chromatophores on the fins. included Gladiogobius ensifer in the key to
Distribution and habitat. Type specimens of species of Japanese gobies. However, their “G.
Gladiogobius brevispinis was collected from ensifer” appears to comprise three species. The
three widely separated localities including the specimen upon which their head illustration was
Ryukyu Islands of Japan, Hainan Island in the based (Akihito et al., 1993: 1012; 2000: 1238;
South China Sea, and Indonesia. The species is 2002: 1238) was correctly identified as G.
certainly more widely distributed and more ensifer, but the specimen used in the line drawing
202 Koichi Shibukawa and Gerald R. Allen
of the entire body (Akihito et al., 1993: 1012, 23337, 1 specimen (female), 25.0mmSL, east
2000: 1238; 2002: 1238) is actually G. brevispi- coast of Makabo Island, Florida Islands,
nis, judging from the pigmented pattern of the Solomon Islands, 0.5m depth, 24 Aug. 1984
head and body, short cheek spine, and the lack of (collected by K. Matsuura); NSMT-P 61522, 1
elongate spines on the first dorsal fin. They also specimen (male), 24.8mmSL, Tanjung Lampu,
noted that G. ensiferis distributed in the Red Sea west coast of Lembeh Island, eastern off Bitung
and Indo-West Pacific region, which suggests of Sulawesi, Indonesia, 1m depth, 13 July 2000
that their “G. ensifer” also contained G. rex, the (collected by K. Shibukawa); NSMT-P 76627, 1
only known species of the genus found in the specimen (female), 34.0mmSL, Iwayama Bay,
Red Sea and Indian Ocean (see also “Remarks” Palau Islands, 1.0m depth, 16 June 1980 (col-
for G. rex). lected by K. Shibukawa); NSMT-P 76628, 3
The typical fleshy crest on the nape is not de- specimens (1 males and 2 females, including 1
veloped in the smallest specimen that was exam- female cleared and stained), 24.3–26.0mmSL,
ined (one of URM-P 36312, 12.5mmSL). Sendakan, west coast of Lembeh Island, eastern
Etymology. The new species is named bre- off Bitung of Sulawesi, Indonesia, 17 July 2000
vispinis (Latin, meaning “short-spine”) in refer- (collected by K. Shibukawa); URM-P 24400, 2
ence to the distinctive preopercle spine, which is specimens (2 females), 26.3–29.6mmSL, Ngth-
the shortest for the genus. The new Japanese ver- an, Babeldaub, Palau Islands, 29 Aug. 1990 (col-
nacular is a combination of the Japanese words lected by T. Yoshino); WAM 32242-001, 2 speci-
“ko” (meaning “small”), “toge” (meaning mens, 30.5–34.9mmSL, Misool Island, Indone-
“spine”) and “haze” (meaning “goby”). sia (collected by G. R. Allen); YCM-P 26084, 2
specimens (1 male and 1 female), 30.1–33.3mm
SL, off Atetsu, Setouchi-cho, Oshima-gun,
Gladiogobius ensiferHerre, 1933 Amami-oshima Island, Kagoshima Prefecture,
(Figs. 1A, 2B, 3B, 4) Japan, 26 Aug. 1991 (collected by Sagami-bay
[Japanese name: Togenaga-haze] Marine Biological-research Club); YCM-P
[English name: Bandit goby]
28509, 1 specimen (male), 30.0mmSL, off Atet-
Gladiogobius ensifer Herre, 1933: 23 [original descrip- su, Setouchi-cho, Oshima-gun, Amami-oshima
tion, type locality: near entrance to Majalibit Inlet, Island, Kagoshima Prefecture, Japan, 31 Aug.
Waigiu ((cid:3)Waigeo Island), Indonesia]; Koumans, 1940: 1992 (collected by Sagami-bay Marine Biologi-
147 [notes on paratypes and cotypes]; Koumans, 1953:
cal-research Club); YCM-P 28555, 1 specimen
32–33 [part: his line drawing of this species in fig. 6
(male), 31.1mmSL, collecting data same with
was based on G. rex (see also remarks under G. rex,
below)]; Hayashi et al., 1981: 5, pl. 1, fig. 100 (Ishiga- YCM-P 28509. In addition, the digital image and
ki-jima Island, Ryukyu Islands, Japan); Akihito et al., x-ray of holotype of Gladiogobius ensifer
1984: 233 (part), pl. 353, fig. D; Akihito et al., 1993: (FMNH 17412, 32.5mmSL, Fig. 4) were exam-
1093 (head illustration showing cephalic sensory sys-
ined (see remarks section, below).
tems), 2000: 1302 (head illustration showing cephalic
Diagnosis. Gladiogobius ensifer is distin-
sensory systems), 2002: 1302 (head illustration show-
guished from other congeners in having the fol-
ing cephalic sensory systems); Masuda and Kobayashi,
1994: 328, fig. 8 (Yaeyama Group of Ryukyu Islands); lowing combination of characters: cheek spine
Myers, 1999: 255, pl. 161, fig. C (Palau, Micronesia); moderately long (its length 21.3–26.7% in head
Kuiter and Tonozuka, 2001: 676, figs. B and C (Bali, length), but not reaching posteriorly to posterior
Indonesia); Shibukawa et al., 2003: 184 (Bitung of Su-
margin of opercular membrane; I, 8–9 (typically
lawesi Island, Indonesia).
I, 9) anal-fin rays; fourth spine of first dorsal fin
Materials examined. Sixteen specimens, greatly elongate, filamentous, longer than preced-
24.3–34.0mmSL: AMS I.24830-001, 2 speci- ing spine; 10–12 (typically 10 or 11) transverse
mens, 28.8–32.5mmSL, Palau Islands; NSMT-P scales from anal-fin origin upward and forward