Bull. Natl. Mus. Nat. Sci., Ser. A, Suppl. 1, pp. 185–205, March 22, 2007

Three New Species of the Shrimp-associated Goby Genus Vanderhorstia(Perciformes: Gobiidae: Gobiinae) from Japan, with
Re-descriptions of Two Related Congeners
Akihisa Iwata1, Koichi Shibukawa2 and Nobuhiro Ohnishi3
1

Division of Southeast Asian Area Studies, Kyoto University, 46 Shimoadachi-cho,
Yoshida, Sakyo-ku, Kyoto 606–8501, Japan
E-mail: iwata@asafas.kyoto-u.ac.jp
2
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
3
Department of Bioenvironmental Design, Faculty of Bioenvironmental Science, Kyoto
Gakuen University, 1–1 Nanjyo Otani, Sogabe-cho, Kameoka City, Kyoto 621–8555, Japan

Abstract Three new species of the shrimp-associated goby genus Vanderhorstia are describedbased on the specimens collected from the southwest coast of Kochi Prefecture, Shikoku, Japan.
Within the genus, these 3 species are assigned to the phenetically close-knit subgroup (named“large-scale group”), characterized by having the combination of 11 or 12 segmented rays on second dorsal fin and 26–45 longitudinal scale count, as well as relatively reduced sensory-papillae
rows below eye. Vanderhorstia hiramatsui sp. nov. (2 specimens, 62.9–65.1 mm SL) differs fromthe other species of this subgroup in having 12 dorsal-fin soft rays (vs. almost always 11 in the latter). Vanderhorstia kizakura sp. nov. (single specimen, 34.9 mm SL) is readily distinguished fromthe other congeners in having separated pelvic fins (i.e., connecting membrane between innermost
pelvic-fin soft rays greatly reduced). Vanderhorstia rapa sp. nov. (single specimen, 50.3 mm SL)
closely resembles V. auropunctata in squamation (e.g., entirely or almost naked predorsal midline,
and naked pectoral-fin base) and meristic counts (33–35 longitudinal scale count, 11–12 transverse
scale count from origin of second dorsal fin backward and downward to anal-fin base), but differsfrom the latter in having a deep trough along posterior margin of eye (vs. undeveloped in the latter), larger jaws (jaw length 50.8% of head length vs. 43.9–44.1%), and first dorsal fin with minuteyellow spots only (vs. narrow yellow lines on distal half). The large-scale group of Vanderhorstia
is re-defined, 2 poorly-known congeners (i.e., V. longimanus and V. puncticeps) are re-described,
and a key to all-known species of this subgroup is also provided.
Key words : Vanderhorstia, new species, Gobiidae, shrimp-associated goby.

Fishes of the Indo-Pacific gobiid genus Vanderhorstia Smith, 1959 are small- to moderatesized marine gobies, symbiotically associatedwith alpheid shrimps (snapping shrimps or pistol
shrimps). The genus currently recognized is characterized by having the following combination ofcharacters (Shibukawa and Suzuki, 2004; present
study): typical “Priolepis Group” type of axial
skeletal features (Birdsong et al., 1988), i.e.,
101626 vertebrae, P-V 3/II II I I 0/9, 2 anal-

fin pterygiophores anterior to first haemal spine,and single epural; wide gill opening, extending
anteriorly to well beyond a vertical line throughposterior preopercular margin; branchiostegal
membranes narrowly attached to isthmus (notforming free rear margin across isthmus); firstgill slit moderately broad, i.e., ventral one-thirdor less of lower arch closed by membrane; noscales on cheek and operculum; relatively longcaudal fin, longer than head; pelvic fins united

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medially by well developed frenum (between
spines) and interradial connecting membrane(between innermost rays) [note. —one of the
new species herein described has almost separated pelvic fin]; vomer and palatine edentate; longitudinal pattern of sensory-papillae rows on
cheek; sensory-papillae row c oblique and moderately long, its terminus close to rows a and b(row a apparently continuous with row c in somespecies); row cp comprising a single papilla; apair of sensory papillae just behind chin (rowf ); cephalic sensory canals well developed, withpores B, C (unpaired), D (unpaired), E, F, G, H,K, L, M, N and O. The other shrimp-goby
genus Ctenogobiops Smith, 1959 resembles Vanderhorstia in sharing most of these features, but
the former is chiefly distinguished from the latterin having relatively shorter caudal fin (subequalor shorter than head in many species) and characteristic bright white spot on pectoral fin [recently
Randall et al. (2003) re-diagnosed Ctenogobiops,
but we could not well separate Ctenogobiops
from Vanderhorstia based only on the charactersthey provided]. Also, Amblyeleotris Bleeker,
1874, another Indo-Pacific shrimp-goby genus
with a pair of sensory papillae just behind chin(Satapoomin and Winterbottom, 2002), differsfrom Vanderhorstia and Ctenogobiops in having
several transverse rows of sensory papillae on
cheek. The following 12 described species are
currently assigned to Vanderhorstia (Shibukawa
and Suzuki, 2004; Winterbottom et al., 2005;
Allen and Randall, 2006): V. ambanoro (Fourmanoir, 1957); V. auropunctata (Tomiyama,1955); V. bella Greenfield and Longenecker,
2005; V. delagoae (Barnard, 1937) (type speciesof Vanderhorstia); V. flavilineata Allen and Munday, 1995; V. longimanus (Weber, 1909); V.
macropteryx (Franz, 1910); V. mertensi Klausewitz, 1974; V. nannai Winterbottom, Iwata andKozawa, 2005; V. nobilis Allen and Randall,
2006; V. ornatissima Smith, 1959; V. papilio
Shibukawa and Suzuki, 2004. Shibukawa andSuzuki (2004) suggested that Vanderhorstia
praealta Lachner and McKinney, 1981 and Vanderhorstia lanceolata Yanagisawa, 1978 ap-

peared to be not placed in the genus; these 2
species should be assigned to the other shrimpassociated goby genus Flabelligobius Smith,1956 or Tomiyamichthys Smith, 1956 (Shibukawa
and Iwata, 2005, in prep.).
Shibukawa and Suzuki (2004) described the
new species Vanderhorstia papilio based on asingle specimen collected from the Ryukyu Islands, Japan, and indicated that it was closed to
two Japan-endemic congeners, V. macropteryx
and V. auropunctata, as well as a single unidentified species from Japan. These 4 species form aphenetically close-knit assemblage, sharing relatively low number of longitudinal scale count(i.e., less than 40) and dorsal- and anal-fin segmented rays (i.e., usually 11 or 12) and reduced
sensory-papillae rows below eye; this subgroup
was called as “large-scale group.” Shibukawa andSuzuki (2004) have overlooked another congener,
V. longimanus, known from Indonesia and Australia, which also belongs to this subgroup.
More recently, Winterbottom et al. (2005) described Vanderhorstia nannai as a new species
from Palau and the Philippines. This species
could be assigned to the large-scale group by
having 11 dorsal/anal segmented rays and characteristic configuration of sensory-papillae rows
on cheek (e.g., row a comprising 3 or 4 widelyseparated papillae), although it has slightly more
longitudinal scale count (41–45) than the othercongeners assigned to this subgroup.
Early in 2006, we had an opportunity to examine 2 specimens of a goby from Tosa Bay, off Pacific coast of Shikoku, Japan, as well as the color
photographs of one of them. Judging from its
characteristic coloration and the other general
physiognomy, the species was identified as Vanderhorstia puncticeps (Deng and Xiong in Xu
et al., 1980) that had been hitherto recognized asa species of Ctenogobius (Xu et al., 1980; Cheng
and Zheng, 1987). These newly-obtained specimens also revealed that the unidentified species
reported by Shibukawa and Suzuki (2004) was
conspecific with V. puncticeps.
In this paper, we describe 3 new species of thelarge-scale group of Vanderhorstia from theNew Species of Vanderhorstia

Japanese waters, re-describe 2 poorly-known
congeners, i.e. V. longimanus and V. puncticeps,
and provided a key to all known species of thelarge-scale group. All new species described inthis paper were already reported by Senou et al.
(2004) as 3 of 8 unidentified species of Vanderhorstia found in Japanese waters; the remaining
5 do not belong to the large-scale group, and,
therefore, are not treated here.
Materials and Methods
Institutional abbreviations for materials examined follow Leviton et al. (1985), except for the
Kanagawa Prefectural Museum, Japan (KPM).
Comparative materials of the species of Vanderhorstia are listed in Shibukawa and Suzuki(2004), except as follows. Vanderhorstia auropunctata: KPM-NI 2791, 1 specimen, female,58.7 mm SL, Suruga Bay, Ose-zaki, Shizuoka
Prefecture, Japan, 53 m depth, 8 Sept. 1996, collected by H. Senou, M. Yanagita and A. Mishiku.
Vanderhorstia nannai: ROM 76552, holotype ofVanderhorstia nannai, 29.5 mm SL, west coast ofBabeldaob, Aimeliik, Palau (7°72.3N, 134°26.0
E), 9.1–15.2 m depth, 19 May 2004, collected by
R. Winterbottom and party; NSMT-P 70085,
paratypes of V. nannai, 10 specimens, 20.8–30.0mm SL, purchased from Oasis Aquarium Store,
Nagoya, Japan, between May 2001 and July
2002.
Measurements were made point-to-point withcalipers under a dissecting microscope to thenearest 0.01 mm. The methods for measurements
follow those of Hubbs and Lagler (1958), withexceptions given below (the snout tip refers to
the mid-anteriormost point of the upper lip): interorbital width is the least width between innermost rims of right and left eyes; jaw length is the
distance between the snout tip and the posteriormost point of lip; head width and depth were
measured at preopercular margin; body depth
was measured at the anal-fin origin; nape width
is the distance between dorsalmost ends of gill
openings; preanal and prepelvic lengths are thedistances from the snout tip to the origin of each187

fin; pectoral-fin length is the length of the longest
ray; pelvic-fin length is the distance between thebase of pelvic-fin spine and the distal tip of thelongest segmented ray; heights of pelvic-fin
frenum and connecting membrane are least
heights; lengths of fin spines and rays are the distances between the base to distal tip of each ray.
Methods for counts follow Akihito (1984), except for the following: longitudinal scale count is
the number of oblique (anterodorsal to posteroventral) scale rows and is taken from just dorsal to the upper attachment of the opercularmembrane posteriorly to the mid-base of caudal
fin; 3 methods of transverse scale count weretaken (see descriptive accounts); circumpeduncular scale count is the number of scales along a
zigzag vertical line through the narrowest pointof the caudal peduncle; gill rakers were counted
on the outer side of first arch, including all rudiments; count of pseudobranchial filaments includes all rudiments. Pectoral- and branched caudal-fin rays were counted and numbered fromdorsal to ventral. Scales (except for predorsal andcircumpeduncular scales), paired-fin rays, gillrakers and pseudobranchial filaments were counted bilaterally. Osteological features were studied
from radiographs. The methods of Akihito(1984) were used in describing the pattern of theinterdigitation 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 follow Akihito (1984) and Miller (1986),
respectively. All fish lengths given are standard
lengths (SL).
Species are arranged in alphabetical order.
Since the species assigned to the “large-scalegroup” of Vanderhorstia share many featureswith one another, we do not include the featuresother than meristic counts and coloration in the“Description” of each species; the other characteristics are found in the accounts of this subgroup, “Diagnosis” of each species account, andTable 1.

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The Large-scale Group of Vanderhorstia
Diagnosis. Fishes of the large-scale group of
Vanderhorstia are distinguished from the othercongeners (hereafter referred as “small-scalegroup”) by having: 11 or 12 segmented rays insecond dorsal fin (vs. 11–16 in the small-scalegroup); 8–12, almost always 11, anal-fin soft rays
(vs. 11–17); 26–45 longitudinal scale count (vs.
44 or more); reduced sensory-papillae row acomprising 3 (or rarely 4) widely-spaced papillae(vs. row a comprises 5 or more papillae); scales
present on predorsal midline in all but 4 species(vs. always absent).
Description. Dorsal-fin rays VI-I, 11–12;
anal-fin rays I, 8–12 (almost always 11); pectoral-fin rays 16–21; pelvic-fin rays I, 5; segmented caudal-fin rays 98, including 6–76–7
branched rays; dorsal unsegmented caudal-fin
rays 6–10; ventral unsegmented caudal-fin rays
5–9; longitudinal scale count 26–45; transverse
scale count from anal-fin origin dorsoanteriorly
to first dorsal-fin base 9–20; transverse scales
count from anal-fin origin dorsoposteriorly tosecond dorsal-fin base 8–16; transverse scalecount from second dorsal-fin origin ventroposteriorly to anal-fin base 7–17; predorsal scales
0–16; circumpeduncular scales 10–12 (typically12); gill rakers 3–59–19; pseudobranchial filaments 6–12; vertebrae 101626; P-V 3/II II I I
0/9; epural 1; anal-fin pterygiophores anterior to
first haemal spine 2; pleural ribs on third to tenthprecaudal vertebrae.
Body moderately slender (body depth 13.0–
17.8% of SL), compressed posteriorly. Headslightly compressed, its width 80.9–94.5% of its
depth. Snout short, its length shorter than eye diameter; snout does not protrude beyond upperlip. Eye dorsolateral, large, its diameter 26.3–
38.1% of head length; interorbital space narrow,
its width narrower than pupil diameter and 3.1–
5.9% of head length. No cutaneous ridge along
dorsal midline of nape. Gape oblique, forming anangle of about 30–50 degrees with body axis.
Lower jaw slightly projecting beyond upper jaw;

posterior end of jaws extending to points between
vertical lines through anterior margin of pupil
and posterior margin of eye. Anterior naris opening at tip of short tube (its height shorter than diameter of its opening); no fleshy flap at tip of anterior naris; posterior naris opening a pore, located at approximately mid-point between anteriornaris and anterior margin of eye. Tongue tip freefrom floor of mouth; anterior margin of tongue
blunt and nearly truncate, without a distinct
notch in the anterior margin. Posteroventral margin of lower lip interrupted at lower-jaw symphysis, except for V. nannai with uninterrupted, freeventral margin across symphysis. Mental flap onchin undeveloped. Gill opening wide, extending
anteriorly well beyond a vertical line throughposterior margin of preopercle; gill membranes
narrowly attach to isthmus; attached gill membranes with no distinct free rear margin acrossisthmus. No fleshy projections on lateral wing ofshoulder girdle. No bony projections along posterior margin of preopercle. Gill rakers on outersurface of ventral arm of first arch well developed, long and thin, finger-like; rakers on outersurface of dorsal arm of first arch short and rudimentary; first gill slit usually well open, ventral
one-seventh to one-third of ventral arm closed bymembrane. Caudal peduncle moderately slender,
its depth 43.9–60.3% of caudal-peduncle length.
First dorsal fin higher than second dorsal fin; firstdorsal fin close to, but not connected to, second
dorsal fin by membrane; all dorsal-fin spines
slender and flexible; all segmented rays of second
dorsal fin branched. Origin of anal fin on a vertical line between bases of first and second segmented rays 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. Caudal fin typically more or lesssymmetrical dorsoventrally, oblong or bifurcate;caudal fin long, its length 115.2–153.4% of headlength. Pectoral fin nearly lanceolate, reaching
posteriorly to points between vertical lines
through bases of first and fifth segmented rays ofsecond dorsal fin, and always extending beyond a
vertical through anus; all pectoral-fin rays

New Species of Vanderhorstia

branched, excluding 1–2 lowermost and/or uppermost simple ray(s). Origin of pelvic fin at, orslightly anterior to, a vertical line through originof first dorsal fin; pelvic fins usually united medially by well developed frenum (between spines)
and connecting membrane (between innermost
rays), except for one species (i.e., V. kizakura)with deeply-concaved connecting membrane;
pelvic frenum moderately thin, with smooth posterior margin; all segmented pelvic-fin rays
branched.
Most of body and caudal-fin base covered by
ctenoid scales with peripheral cteni; ctenoidscale area extending anteriorly to a vertical line
through middle or posterior part of base of firstdorsal fin; cycloid scales on nape (when present),
pectoral-fin base (when present), belly, prepelvic
region, and small areas on anterodorsal part ofbody and behind and below pelvic-fin base; pectoral-fin base covered by cycloid scales in all but
3 species (i.e., V. auropunctata, V. puncticeps andV. rapa); head always naked.
Teeth in both jaws unicuspid, slender, more orless inwardly curved; upper jaw with 3–4 rows ofteeth anteriorly, narrowing to 1–2 rows posteriorly; teeth on middle row(s) of upper jaw smaller
than teeth in outermost and/or innermost rows;
1–3 strongly incurved, enlarged teeth (sometimes
stout and canine-like) anteriorly in innermost row
of upper jaw; lower jaw with 4 rows of teeth anteriorly, narrowing to single row posteriorly; 2–4
stout, enlarged canine-like teeth medially in innermost row of lower jaw; other teeth on lower
jaw subequal in size; no teeth on vomer or palatine. Anterior tip of vomer enlarged in a bluntbilobed process, projecting downward behindsymphysis of upper jaw.
Anterior oculoscapular-canal pores B, C (unpaired), D (unpaired), E, F, G and H [note. —pore G of left side of holotype of Vanderhorstia
kizakura absent]; posterior oculoscapular-canalpores K and L; preopercular-canal pores M, N
and O; right and left sides of anterior oculoscapular canals fused medially in interorbital
space; ventralmost pore of preopercular canal (pore O) opening at point slightly below a hori-189

zontal line through posterior end of sensorypapillae row d. All cephalic sensory-papillae
rows uniserial or comprising a single papilla, notforming multiple lines or aggregations; relatively
reduced longitudinal pattern (could be expressed
as reduced transverse pattern, following Winterbottom and Burridge, 1993a, b) of sensory papillae rows on cheek; row a short and reduced, comprising 3 (rarely 4) widely-spaced sensory papillae on both sides; row b broadly interrupted at
midway, and extending from around ventralmost
part of row a to posterior margin of preopercle;
row c comprising 4 or 5 papillae and relatively
short, its posterior end close to anterior ends ofrows a and b; row cp comprising a single papilla;row d not reaching to, or extends slightly posterior to, a vertical line through row cp; each side ofrow f a single papilla (namely, a pair of papillae
just behind chin); row n comprising a single
papillae; row s2 comprising a single papilla; row
x1 well separated from row x2. Sensory papillae
on midlateral body form uniserial vertical rows,
each row short and restrictedly found on a single
scale; 3 radiating rows of sensory papillae on
caudal fin, each along the fourth, seventh andtenth branched caudal-fin rays; all 3 sensory
papillae rows on caudal fin extend from posterior
margin of scaled area to near distal end of eachray.
Remarks. According to Allen and Munday(1995), Vanderhorstia flabilineata has both 11
segmented rays on second dorsal and anal fins,
likewise the species of the large-scale group;
other species of the “small-scale group” has 14
or more segmented rays on these fins. However,V. flabilineata also has “approximately 50–55(some scales missing)” longitudinal scales andthe sensory-papillae row a appears to comprise 5
sensory papillae (Allen and Munday, 1995:104,
fig. 8), and, thus, we do not assign it to the largescale group tentatively. The large-scale groupherein defined is only the artificially-compiled
assemblage, not characterized by any obvious
synapomorphies supporting its monophyly. In
order to resolve the phylogenetic implication ofthis subgroup, comprehensive study on Am-

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A. Iwata, K. Shibukawa and N. Ohnishi

blyeleotris, Ctenogobiops and Vanderhorstia isneeded (see also “Remarks” of V. kizakura).
Because of the difficulty in collecting correlated in their habits, there are very few museum
specimens of this subgroup. We describe 3 new
species (i.e., V. hiramatsui, V. kizakura and V.
rapa) based only on 1–2 specimens of eachspecies, therefore. Nevertheless, we are convinced that all of these are distinct species, because, as well as the morphological evidence
based on the voucher specimens, we have had theopportunities to examine many excellent underwater photographs of more than one individual ofeach species; some of those images are appearedin the recently-published pictorial books (e.g.,
Hayashi and Shiratori, 2003; Senou et al., 2004)
and the “Image Database of Fishes” in the KPM(also available from the website “FishPix” of the
National Museum of Nature and Science:
http://fishpix.kahaku.go.jp/fishimage-e).
Included species. The following 9 species
are assigned to the large-scale group of Vanderhorstia: V. auropunctata; V. hiramatsui (newspecies); V. kizakura (new species); V. longi-

manus; V. macropteryx; V. nannai; V. papilio; V.
puncticeps; V. rapa (new species). Five species ofthis subgroup (i.e., V. auropunctata, V. hiramatsui,
V. kizakura, V. macropteryx and V. rapa) are hitherto known only from Japanese waters, whereas
the other 4 are found in the other areas of theWest Pacific, as follows: V. longimanus, reported
from Indonesia and Australia; V. nannai, known
from Palau and the Philippines; V. papilio, described based on a single specimen from Japanese waters, and the underwater photographs ofgoby that appear to be identical with V. papillio
were reported from Bali and Sulawesi, Indonesia
by Kuiter and Tonozuka (2001: 633, misidentified as Psilogobius prolatus); V. puncticeps, originally described from Zhejiang Province of China,
and newly recorded here from Japanese waters.
Allen et al. (2003: 312) noted V. macropteryx is
distributed in “Philippines to S. Japan,” but the
species in the underwater photograph given bythem as V. macropteryx is apparently misidentification of V. papilio; we did not locate any certain
evidence suggesting that V. macropteryx is also
distributed outside of Japan.

Key to Species of the Large-scale Group of Vanderhorstia
1a.

Pelvic fins separated, with very low connecting membrane between innermost rays (height ofconnecting membrane 23.3% of length of fifth pelvic fin soft ray); eye large, its diameter
38.1% of head length; 5 narrow dusky bars on head and body, anterior 4 of which diagonalrather than vertical (Shikoku and Ryukyu Islands, Japan)              V. kizakura sp. nov.
1b. Pelvic fins united medially via well-developed connecting membrane between innermost rays(length of connecting membrane 81.1–98.2% of length of fifth pelvic fin soft ray); eye diameter
24.3–37.2% of head length; dusky bars present or absent (if present, 2 bars between anal andsecond dorsal fins almost vertical rather than diagonal)                               2
2a. Predorsal midline entirely or almost naked; pectoral-fin base naked                     3
2b. Nape, including predorsal midline, broadly scaled (except for V. longimanus with only a single
predorsal scale); pectoral-fin base scaled                                          6
3a.

Fifth and ninth branched caudal-fin rays greatly elongate, forming a distinctly bifurcate caudal
fin; lower lip with uninterrupted, free posteroventral margin across dentary symphysis; gape
well oblique, forming an angle of about 50 degrees with body axis; jaw relatively small (length
10.4–13.2% of SL), extending posteriorly to a vertical line through anterior margin of pupil;more than 40 longitudnal scale count; 16–18 pectoral-fin rays; 3–416–19 gill rakers on outer

New Species of Vanderhorstia

191

surface of first arch; 2 rows of orange spots on upper half of body, those on midlateral bodysurrounded by light blue (Palau and Philippines)                             V. nannai
3b. Caudal fin rounded or lanceolate; posteroventral margin of lower lip interrupted at lower-jaw
symphysis; gape moderately oblique, forming an angle of about 30 degrees with body axis;
jaws moderate to relatively large (length 12.5–14.4% of SL), extending posteriorly to a verticalline through middle or posterior margin of eye; 26–35 longitudinal scale count; 20–21 pectoralfin rays; 3–59–11 gill rakers on outer surface of first arch; color not as above            4
4a.

Longitudinal scale count 26–27; 8 transverse scale count from origin of second dorsal fin downward and backward to anal-fin base; when freshly collected, distal margin of first dorsal fin red;a small red spot on operculum (Sagami Bay and Tosa Bay of Japan and Zheijiang Province ofChina) .                                                           V. puncticeps
4b. Longitudinal scale count 32–35; 10–12 transverse scale count from origin of second dorsal fin
downward and backward to anal-fin base; no red markings on first dorsal fin and operculum55a.

Trough along posterior margin of eyes less developed, shallow; jaw moderate in size
(43.9–44.1% of head length), extending posteriorly to a vertical through posterior margin of
pupil (Fig. 1A); fourth spine of first dorsal fin longest, subequal or slightly longer than third
spine (fourth spine 102.5–103.9% of third spine in length); sensory-papillae row c comprising
5 papillae; yellow markings on distal part of first dorsal fin vertically elongated, forming shortlines; second dorsal, anal and caudal fins with a yellow submarginal line; a black blotch sometimes present on first dorsal fin between fourth and fifth or sixth spines (Sagami Bay and Izu Islands, off southern coast of middle of Honshu, Japan)                    V. auropunctata
5b. Deep trough along posterior margin of eyes; jaw relatively large (50.8% of head length), extending posteriorly to a vertical through posterior margin of eye (Fig. 1E); third spine of firstdorsal fin longest, longer than fourth spine (fourth spine 81.0% of third spine in length); row ccomprising 4 sensory papillae; all yellow markings on first dorsal fin more or less circular,
spot-like; second dorsal, anal and caudal fins without yellow submarginal lines; no distinct
black spot on first dorsal fin (Kashiwa-jima Island, off southwest coast of Shikoku, Japan)                                                               V. rapa sp. nov.

6a.

Nape largely naked, with only a single predorsal scale; anterior 3 spines of first dorsal fin greatly elongate and filamentous; body slender, depth 13.0% of SL (Ceram of Indonesia and Western Australia)                                                     V. longimanus
6b. Nape broadly scaled, with 7–18 predorsal scales rows; spines of first dorsal fin not so elongate
and usually non-filamentous (distal tip of third spine forming a short filament in V. papilio);
body less slender, depth 16.8% or more of SL .                                     7
7a.

Fifth and ninth branched caudal-fin rays longer than other rays, greatly elongate and filamentous, forming a distinctly bifurcate caudal fin; 27 longitudinal scale count (Western Pacific)                                                                    V. papilio
7b. Caudal fin more or less rounded without distinctly modified filamentous rays; 30–39 longitudinal scale count                                                               8
8a.

Second dorsal fin with I, 11 rays; no elongate and filamentous spines on first dorsal fin; trans-

192

8b.

A. Iwata, K. Shibukawa and N. Ohnishiverse scale count from origin of second dorsal fin downward and backward to anal-fin base
10–12; no distinct dusky saddle on nape; longitudinal yellow line across middle of cheek; orange line along dorsal margin of upper jaw (off Pacific coasts of southern part of Honshu,
Shikoku and Kyushu, Japan)                                         V. macropteryx
Second dorsal fin with I, 12 rays; third spine of first dorsal fin slightly elongate and filamentous;transverse scale count from origin of second dorsal fin downward and backward to anal-finbase 16–17; black saddle on nape extending ventrally to branchiostegal membrane, obviously
broadened and vivid at operculum; yellow spots only on cheek (i.e., no longitudinal lines); nooran line along dorsal margin of upper jaw (Izu Islands and southwest coast of Shikoku, Japan)                                                          V. hiramatsui sp. nov.

Vanderhorstia hiramatsui sp. nov.
(New Japanese name: Kuroerikanoko-haze)(Figs. 1C, 3 and 4A–B; Table 1)

Vanderhorstia sp. 5: Senou et al., 2004: 365 (underwaterphotograph and brief description; Izu-oshima Island
and Kochi Prefecture of Shikoku, Japan).

Fig. 1. Heads of six species of Vanderhorstia, showing cephalic sensory canal pores (indicated by roman uppercase letters, except for AN and PN) and papillae (indicated by roman lowercase letters). A: Vanderhorstia auropunctata, KPM-NI2791, female, 58.7 mm SL; B: Vanderhorstia kizakura sp. nov., NSMT-P 73010, holotype, female, 34.9 mm SL; C: Vanderhorstia hiramatsui sp. nov., NSMT-P 73121, paratype, female, 62.9 mm
SL; D: Vanderhorstia puncticeps, BSKU 77244, male, 30.3 mm SL; E: Vanderhorstia rapa sp. nov., NSMT-P
73119, holotype, female, 50.3 mm SL. AN and PN, anterior and posterior nostrils, respectively. Arrows show
position where gill membrane attached to isthmus. In the holotype of V. kizakura, sensory-canal pore G andsensory-papillae row z could not be examined, and, in this illustration (B), reproduced based on those of theright side. Bars indicate 3 mm. Drawn by K. Shibukawa.

New Species of Vanderhorstia
Holotype. NSMT-P 73120, 65.1 mm SL, male, Tsutome-zaki Point, Issai, Ostuki, Kochi Pref., Shikoku,
Japan, 41 m depth, 2 May 1998, collected by W. Hiramatsu.
Paratype. NSMT-P 73121, 62.9 mm SL, female,
Tsutome-zaki Point, Issai, Otsuki, Kochi Prefecture,
Shikoku, Japan, 42 m depth, 3 May 1998, collected by W.
Hiramatsu.
Image database of fishes (see “Remarks” of “Thelarge-scale group of Vanderhorstia”). KPM-NR 61047,
Kashiwa-jima Island, Kochi Prefecture, Shikoku, Japan,
20 May 2001, photographed by Y. Miyamoto.

Diagnosis. Vanderhorstia hiramatsui is distinguished from the congeners in having the following combination of characters: 12 segmented
rays on second dorsal fin; 38–39 longitudinalscale count; 15–16 predorsal scales; 16–17 transverse scale count from origin of second dorsal finbackward and downward to anal-fin base; pectoral-fin base broadly covered by cycloid scales;connecting membrane between innermost rays
well developed; trough along posterior margin ofeye well developed; operculum almost entirely
blackened; cheek and occipital region scatteredwith minute yellow spots (and no yellow lines).
Description. In the following description,
the counts of holotype are asterisked, and the frequency of each count is given in the parentheses
following relevant count. Dorsal-fin rays VI-I,
12* (2); anal-fin rays I, 11* (2); pectoral-fin rays
18 (1) or 20* (3); pelvic-fin rays I, 5* (4); segmented caudal-fin rays 98* (2), including 7
7* (2) branched rays; dorsal unsegmented caudal-fin rays 9* (2); ventral unsegmented caudalfin rays 8* (2); longitudinal scale count 38 (2) or
39* (2); transverse scale count from anal-fin origin dorsoanteriorly to first dorsal-fin base 18 (1),19* (2) or 20* (1); transverse scale count fromanal-fin origin dorsoposteriorly to second dorsalfin base 15* (2) or 16* (1); transverse scale countfrom second dorsal-fin origin ventroposteriorly toanal-fin base 16* (3) or 17 (1); predorsal scales
15 (1) or 16* (1); circumpeduncular scales 12*(2); gill rakers 310* (1), 49 (1), 411* (1) or
510 (1); pseudobranchial filaments 11* (3) or
12 (1).
Color when alive [based on the underwaterphotograph found in Senou et al. (2004: 365)].

193

Ground color of head and body pale gray, a little
tinged with blue, darkener dorsally; dorsal surface of head and body tinged with light green; 5vertical dark-brown bars on head and body, firstbar (darkest and more completely developed)from nape downward to ventral part of operculum, second bar from posterior part of first dorsal-fin base downward to belly, third bar from anterior part of second dorsal-fin base (between
fourth and seventh rays) downward to just above
anterior part of anal-fin base, fourth bar at anteriormost part of caudal peduncle, and the fifth andposteriormost one at caudal fin base; pupil edged
narrowly by yellow; iris dark bronze to black
dorsally, yellow to yellowish gray ventrally;
cheek, anterodorsal part of operculum, occipitalregion and body scattered with minute yellow
spots, those on midlateral part of tail of bodycontinuous forming irregular longitudinal line;
first dorsal fin translucent, a little tinged with yellowish gray or blue, scattered with minute yellow
spots; second dorsal fin subtranslucent, a little
tinged with blue, scattered with yellow spots(slightly larger than those on first dorsal fin),with a yellow submarginal line; caudal fintranslucent, slightly tinged with blue, with elongate yellow spots and lines on its dorsal and ventral part, respectively; pectoral fin largely transparent; pelvic fin pale blue with yellow lines.
Color in alcohol. Similar to live coloration,
except as follows: ground color of head and body
pale yellow or pale brown, becoming slightly
darkener dorsally; all yellow spots on head, bodyand fins pale; bluish areas of fins turn to darkbrown; iris entirely blackish.
Distribution and habitat. Vanderhorstia hiramatsui is described based on 2 specimens collected from the southwest coast of Shikoku,
Japan. Senou et al. (2004) reported that thisspecies (as Vanderhorstia sp. 5) distributed in
Izu-oshima Island (Izu Islands, off pacific coastof middle of Honshu) and Kashiwa-jima Island(off southwest coast of Shikoku).
Etymology. The new species is named hiramatsui in honor of W. Hiramatsu, who collected
the type specimens of this species.

194

A. Iwata, K. Shibukawa and N. Ohnishi

Remarks. Vanderhorstia hiramatsui is readily distinguished from the other members of the“large-scale group” in having 12 segmented rays
on second dorsal fin (vs. almost always 11 in thelatter, i.e., other than V. hiramatsui, 12 segmented-rays condition is examined only in a single
specimen of V. puncticeps) and almost entirely
blackened operculum (vs. not such color).
One or 2 caudal fin rays are slightly elongatedin the type specimens of Vanderhorstia hiramatsui, but the length of these rays is shorter in thisspecies from those in V. papilio: slightly prolonged, lengths of these rays 102.0–103.7% ofmiddle caudal-fin ray (vs. greatly prolonged,
lengths of these rays 143.0–146.9% of middle
caudal-fin ray in V. papilio); 3 or more branches

as for the neighboring branched rays (vs. dichotomous).
Vanderhorstia kizakura sp. nov.
(New Japanese name: Kizakura-haze)(Figs. 1B, 2A and 4C–E; Table 1)
Gobiidae, indet. gen. & sp.: Anker, 2000: 5, fig. 5 (underwater photograph; Kochi Prefecture, Japan).
Vanderhorstia sp.: Hayashi and Shiratori, 2003: 160,
image No. 313 (underwater photograph and brief description; Kume-jima Island, Japan).
Vanderhorstia sp. 8.: Senou et al., 2004: 367 (underwaterphotograph and brief description; Kochi Prefecture of
Shikoku, Amami-oshima Island, Okinawa-jima Island
and Kume-jima Island, Japan).
Holotype. NSMT-P 73010, 34.9 mm SL, female,Fig. 2. Freshly collected specimens of four species of Vanderhorstia. A: Vanderhorstia kizakura sp. nov.,
NSMT-P 73010, holotype, female, 34.9 mm SL, Kashiwa-jima Island, Kochi Prefecture, Japan, photographedby A. Iwata; B: Vanderhorstia puncticeps, BSKU 77244, male, 30.3 mm SL, Tosa Bay, off Kochi Pref., Japan,
photographed by E. Katayama; C: Vanderhorstia rapa sp. nov., NSMT-P 73119, holotype, female, 50.3 mm
SL, Kashiwa-jima Island, Kochi Prefecture, Japan, photographed by A. Iwata.

New Species of Vanderhorstia
Kashiwa-jima Island, off southwest coast of Shikoku,
Japan, 30 m depth, 4 Aug. 1993 (kept in aquarium aftercollection, and fixed on 30 Nov. 1993), collected by A.
Iwata and N. Ohnishi.
Image database of fishes (see “Remarks” of “Thelarge-scale group of Vanderhorstia”). KPM-NR 16765,
Kashiwa-jima Island, Kochi Prefecture, Shikoku, Japan,
37 m depth, 28 June 1998, photographed by K. Yamazaki;
KPM-NR 27726, Kashiwa-jima Island, Kochi Prefecture,
Shikoku, Japan, 38 m depth, 14 Nov. 1998, photographedby K. Yamazaki; KPM-NR 28876, Kume Island, Okinawa
Group of Ryukyu Islands, Japan, 60 m depth, 6 June
1999, photographed by A. Mishiku; KPM-NR 28877,
Kume Island, Okinawa Group of Ryukyu Islands, Japan,
60 m depth, 6 June 1999, photographed by A. Mishiku;
KPM-NR 36004, Seragaki, Kunigami, Okinawa-jima I.,
Okinawa Group of Ryukyu Islands, Japan, 48 m depth, 8
Dec. 1999, photographed by M. Takata; KPM-NR 38868,
Kashiwa-jima Island, Kochi Prefecture, Shikoku, Japan,
40 m depth, 1 Sep. 2000, photographed by A. Okumura;
KPM-NR 63099, Kashiwa-jima Island, Kochi Prefecture,
Shikoku, Japan, May 2001, photographed by A. Okumura; KPM-NR 63428, Kashiwa-jima Island, Kochi Prefecture, Shikoku, Japan, 20 Nov. 2001, photographed by K.
Matsuno; KPM-NR 63429, Kashiwa-jima Island, Kochi
Pref., Shikoku, Japan, 15 Oct. 2001, photographed by K.
Matsuno.

Diagnosis. The new species Vanderhorstia
kizakura is unique within the genus in having
greatly reduced connecting membrane between
innermost pelvic-fin rays, i.e., its height 23.3% oflength of fifth pelvic-fin soft rays. The species isalso distinguished from the congeners in having
the following combination of characters: 11 segmented rays on second dorsal fin; 33–34 longitudinal scale count; 11 predorsal scales; 11–12
transverse scale count from origin of second dorsal fin backward and downward to anal-fin base;pectoral-fin base with cycloid scales; third spineof first dorsal fin longest, slightly elongate andfilamentous; ninth segmented caudal-fin ray
slightly elongate, forming a pin-tailed caudal fin;
5 narrow dusky bars on head and body, anterior 4
diagonal rather than vertical; some short diagonal yellow lines on operculum.
Description. In the following description,
the values where we take bilaterally are separated
by a slash, the first value representing the left
count. Dorsal-fin rays VI-I, 11; anal-fin rays I,

195

11; pectoral-fin rays 19/20; pelvic-fin rays I, 5;
segmented caudal-fin rays 98, including 77
branched rays; dorsal unsegmented caudal-fin
rays 8; ventral unsegmented caudal-fin rays 8;
longitudinal scale count 33/34; transverse scalecount from anal-fin origin dorsoanteriorly to firstdorsal-fin base 17/18; transverse scale countfrom anal-fin origin dorsoposteriorly to second
dorsal-fin base 13/13; transverse scale countfrom second dorsal-fin origin ventroposteriorly toanal-fin base 11/11; predorsal scales 11; circumpeduncular scales 12; gill rakers 310/310;
pseudobranchial filaments 10/9.
Color when alive or fresh [based on Fig. 2B
and the underwater photographs found in, e.g.,
Senou et al. (2004: 367)]. Ground color ofhead and body pale, slightly grayish dorsally;
belly pale white; five narrow dusky, vertical orslightly diagonal bars on head and body, posterior 4 of which tinged with yellow; first bar fromnape downward to operculum, second bar fromposterior part of base of first dorsal fin downward
to middle of belly, third bar from middle of baseof second dorsal fin (between third and seventhrays) downward to just above anterior part ofanal-fin base (between first and third soft rays),
fourth bar from anteriormost part of dorsal surface of caudal peduncle downward to just above
posteriormost part of anal-fin base (between
tenth and eleventh soft rays), and fifth and posteriormost bar at caudal-fin base; horizontallyelongate triangular yellow patch beneath eye; ashort, narrow bright sky-blue line along posteroventral margin of eye; occipital region withshort diagonal yellow lines and small spots;
cheek scattered with many various-sized (allsmaller than pupil) yellow spots; operculum with
3 short, almost vertical yellow lines, dorsalmostof which extending dorsally to nape; numerous
minute dust-like yellow dots scattered on dorsal
part of body; first dorsal fin translucent andslightly whitish, with narrow yellow lines alongspines; second dorsal fin translucent and slightly
whitish, with 2 vague yellow basal blotches andnarrow yellow submarginal line; short yellow
lines along middle of spine and rays of second

29.0
16.0
18.0
6.6
8.9
1.7
12.5
14.5
17.5
13.1
34.7
34.3
61.1
19.2
11.6
20.9
26.9
23.3
30.5
27.5
35.5
17.1
21.0
24.1
18.0
13.1
14.9
24.0
9.0
11.5
22.8
8.7
13.6
26.2
25.7

% of SL
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
D1 baseD2 baseA baseP1 length
P2 length
C length
Length of 1st spine of D1
Length of 2nd spine of D1
Length of 3rd spine of D1
Length of 4th spine of D1
Length of D2 spine
Length of 1st ray of D2
Length of longest ray of D2
Length of A spine
Length of 1st ray of A
Length of longest ray of A
Length of P2 spine
Length of 1st ray of P2
Length of 4th ray of P2
Length of 5th ray of P2
29.7
15.9
18.3
7.3
8.9
1.7
12.9
14.5
17.4
13.0
36.2
33.9
60.5
19.7
11.6
21.7
27.4
22.6
30.5
27.0
36.3
19.6
23.2
28.9
20.4
13.7
15.2
21.5
8.6
11.8
20.6
8.1
14.0
26.2
25.2

62.9

NSMT-P 73121
Female

26.1
15.5
17.9
5.9
9.9
1.5
11.7
13.2
17.8
15.4
33.7
30.3
56.9
20.6
11.1
21.1
28.4
24.7
29.1
25.3
35.8
17.0
19.2
19.6
15.0
10.1
13.8
18.9
8.0
10.2
20.2
8.6
11.4
23.1
23.9

34.9

NSMT-P 73010
Female

26.9
14.5
16.0
5.4
9.5
1.2
10.5
12.5
15.1
13.0
29.4
35.1
55.1
21.0
9.9
19.6
26.4
26.3
26.2
19.0
37.2
24.4
24.5
24.5
21.7
9.9
12.8
22.2
8.4
10.5
21.4
6.7
8.9
16.6
17.9

39.9

NSMT-P R798
Male

25.9
12.6
15.6
5.6
7.7
1.3
10.4
12.7
14.9
13.1
32.9
32.6
55.4
20.8
9.1
19.2
30.0
26.3
26.5
19.6
39.7
22.8
22.1
21.0
19.0
10.3
12.8
20.9
7.3
10.7
—
5.8
10.0
18.2
18.1

33.3

BSKU 77244
Male

V. puncticeps

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

65.1

Standard length (SL)NSMT-P 73120
Male

V. kizakura sp. nov.

Proportional measurements of five species of Vanderhorstia.

V. hiramatsui sp. nov.

Table 1.

28.7
13.7
14.5
5.1
10.7
1.2
9.4
12.4
13.0
9.1
34.2
33.9
56.4
19.8
9.4
21.9
26.9
21.9
25.8
22.9
37.2
24.1
27.3
37.0
23.2
—
—
—
—
—
—
7.3
11.7
21.0
21.7

25.2

ZMA 110.978
FemaleV. longimanus

28.4
15.4
17.1
5.8
8.6
1.4
10.3
14.4
15.9
13.8
34.2
32.5
58.7
20.6
10.7
20.8
26.1
22.6
27.4
21.8
33.3
21.4
23.1
23.1
18.7
11.7
14.1
21.2
7.3
9.6
19.9
6.9
10.8
20.2
20.3

50.3

NSMT-P 73119
FemaleV. rapa sp. nov.

196
A. Iwata, K. Shibukawa and N. Ohnishi

New Species of Vanderhorstia

197

Fig. 3. Preserved specimens of Vanderhorstia hiramatsui sp. nov. A) NSMT-P 73120, holotype, male, 65.1 mm
SL, Kashiwa-jima Island, Kochi Prefecture, Japan; B) NSMT-P 73121, paratype, female, 62.9 mm SL, Kashiwa-jima Island, Kochi Prefecture, Japan. Photographed by K. Shibukawa.

dorsal fin; anal fin slightly whitish, with a yellow
submarginal line margined by blue dorsally andventrally; spine and soft anal-fin rays tinged withyellow; caudal fin subtranslucent and slightly
whitish; submarginal yellow line margined byblue dorsally and ventrally on dorsal and ventral
part of caudal fin; distal part of branched caudalfin rays tinged with yellow distally; pectoral fintranslucent; pelvic fin pale white.
Color in alcohol. Similar to live coloration,
except as follows: ground color of head and body
pale yellow or pale brown, becoming darkener
dorsally; all yellow spots on head, body and finsturn to pale; bluish areas of fins turn to darkbrown; iris entirely blackish.
Distribution and habitat. The new species
Vanderhorstia kizakura is described based on asingle specimen from Kashiwa-jima Island, offsouthwest coast of Shikoku, Japan. Senou et al.
(2004) reported that this species was known from
Kochi Prefecture of Shikoku and the Ryukyu Islands (Amami-oshima Island, Okinawa-jima Island and Kume-jima Island). It is found in sandy
gentle slope below coral-reef drop-off (45 m or
more depths), and symbiotically associates withthe alpheid shrimps (Anker, 2000; Senou et al.,2004).
Etymology. The new specific name, kizakura,is the Japanese vernacular name for a variety of
the cherry tree with turmeric-colored blossoms,
in reference to the bright yellow dots and lines
scattered on head and body of this species. The
name should be treated as noun in apposition.
Remarks. When this new species was discovered, it was first considered to be an undescribed species of Amblyeleotris because of its
general physiognomy, e.g., almost separated
pelvic fins and characteristic banded pattern onhead and body. Subsequent our examination reveals that it has longitudinal pattern of sensorypapillae rows on cheek, suggesting that it belongs to Vanderhorstia (vs. several transverserows of sensory papillae below eye in Amblyeleotris).
Amblyeleotris is distinguished from Vanderhorstia based chiefly on this discrepancy in theconfiguration of sensory-papillae rows below
eye. However, each transverse row of sensory
papillae on cheek appears to correspond to eachsensory papillae of rows a, c, and cp in the large-

198

A. Iwata, K. Shibukawa and N. Ohnishi

Fig. 4. Underwater photographs of three species of Vanderhorstia. A: Vanderhorstia hiramatsui, ca. 8 cm TL,
Tsutome-zaki Point, Kochi Prefecture, Japan, 44 m depth, 4 Oct. 1998; B: Vanderhorstia hiramatsui, ca. 7 cm
TL (total length), Tsutome-zaki Point, Kochi Prefecture, Japan, 39 m depth, 26 Apr. 1998; C: Vanderhorstia
kizakura, ca. 4 cm TL, Kashiwa-jima Island, Kochi Prefecture, Japan, 35 m depth, 29 Apr. 1996; D: Vanderhorstia kizakura, ca. 5 cm TL, Kashiwa-jima Island, Kochi Prefecture, Japan, 35 m depth, 23 Nov. 1995; E:
Vanderhorstia kizakura, ca. 5 cm TL, Kashiwa-jima Island, Kochi Prefecture, Japan, 35 m depth, 2 June 1996;
F: Vanderhorstia rapa, ca. 4.5 cm TL, Kashiwa-jima Island, Kochi Prefecture, Japan, 36 m depth, 20 July
1997; G: Vanderhorstia rapa, ca. 4 cm TL, Kashiwa-jima Island, Kochi Prefecture, Japan, 35 m depth, 29
Apr. 1996; H: Vanderhorstia rapa, 5 cm TL, Izu-oshima Island, Izu Islands, Japan, 37 m, 14 Nov. 2005. Photographed by T. Hirata (A–G) and Y. Miyamoto (H).

New Species of Vanderhorstia

scale group of Vanderhorstia and its putative relative, Ctenogobiops; in addition, number of sensory papillae in each transverse row varies between the species of Amblyeleotris (see, e.g., Akihito et al., 2002). Similar proliferation/reductionof sensory papillae is found in various gobiid
genera with essentially “longitudinal pattern”
of sensory-papillae rows, e.g., Acentrogobius,
Favonigobius, Glossogobius, Oligolepis, andRhinogobius, as intrageneric variation (e.g.,
Hoese, 1983; Akihito et al., 2002). To ascertain
the interrelationships of the close-knit group of
the genera Amblyeleotris, Ctenogobiops and Vanderhorstia is beyond the scope of this study, andherein we tentatively follow the generic definition currently recognized. Discovery of Vanderhorstia kizakura, having the intermediate appearance between Vanderhorstia and Amblyeleotris,suggests that a reevaluation of their statuses isrequired.
Vanderhorstia longimanus (Weber, 1909)(Fig. 5; Table 1)
Gobius (Oxyurichthys) longimanus Weber, 1909: 154(original description); Weber, 1913: 478, fig. 97 (description and line drawing).
Oxyurichthys longimanus: Koumans, 1953: 41–42, fig. 10[description and line drawing (after Weber, 1913)].
?Vanderhorstia longimanus: Hutchins, 2001: 44 (checklist; Western Australia).
Materials examined. ZMA 110.978, holotype of
Gobius (Oxyurichthys) longimanus, female, 25.2 mm SL,
Ceram, Indonesia (2°28.52S, 131°3.30E), 118 m depth,
22 Aug. 1899, collected by M. Weber.

Diagnosis. Vanderhorstia longimanus is distinguished from the congeners in having the following combination of characters: 11 segmented
rays on second dorsal fin; 30–31 longitudinalscale count; single predorsal scale; 8–9 transverse scale count from origin of second dorsal finbackward and downward to anal-fin base; pectoral-fin base scaled; anterior 3 spines of firstdorsal fin greatly prolonged and filamentous, extending posteriorly to a little beyond base of seventh segmented ray of second dorsal fin; caudal

199

fin long, lanceolate; in preservative, cheek, operculum and nape with several pale spots; aseries of 4 dusky blotches on side of body.
Description. In the following description,
the values where we take bilaterally are separated
by a slash, the first value representing the left
count. Dorsal-fin rays VI-I, 11; anal-fin rays I,
11; pectoral-fin rays 19/19; pelvic-fin rays I, 5/I,
5; longitudinal scale count 31/30; transverse
scale count from anal-fin origin dorsoanteriorly
to first dorsal-fin base 10/9; transverse scalecount from anal-fin origin dorsoposteriorly tosecond dorsal-fin base 9/8; transverse scale countfrom second dorsal-fin origin ventroposteriorly toanal-fin base 8/9; predorsal scale 1; circumpeduncular scales 12; gill rakers 38; pseudobranchial filaments 8.
Color in alcohol. Head and body pale
brown; dorsal, anal, caudal and pectoral fins pale;
all pigmented patterns already faded, except for
middle of pelvic fins dusky. Color in the line
drawing of the holotype provided by Weber(1913) and Koumans (1953) is as follows: a series of 4 dusky blotches on midlateral body; firstdorsal fin entirely covered by numerous dense
melanophores; cheek and upper part of operculum covered by numerous dense melanophores,
with several small pale spots (Koumans noted“some pearl like spot on cheek, opercle andnape”).
Distribution and habitat. The holotype of
this species was collected from the Ceram Sea,
Indonesia, at the depth of 118 m. It had been
known only by the holotype until recently; although we have not examined the voucher specimen(s), Hutchins (2001) reported it from Western Australia.
Remarks. Weber (1909) described Gobius(Oxyurichthys) longimanus based on a single
specimen from the Ceram Sea. Later, Koumans(1953) listed it as Oxyurichthys longimanus.
Koumans correctly accounted that this species
had “In interorbital anteriorly and posteriorly inmedian line an open pore”; as confirmed in thisstudy, this means that it has unpaired pores C and
D, indicating it belongs to the gobiid subfamily

200

A. Iwata, K. Shibukawa and N. Ohnishi

Fig. 5. Vanderhorstia longimanus. Top) ZMA 110.978, holotype of Gobius (Oxyurichthys) longimanus, female,25.2 mm SL, Ceram, Indonesia, photographed by K. Shibukawa; bottom) line drawing of the holotype [after
Weber, 1913 (horizontally reversed)].

Gobiinae (sensu Pezold, 1993). Oxyurichthys
Bleeker, 1857, is the gobionelline genus (sensu
Pezold, 1993), and has a paired pore C (Akihito
et al., 1984, 1993, 2002; Pezold, 1991, 1993,
1998).
On the contrary, there are several differences
between the description made by the previous authors and the actual features of the holotype in
the number of scales [e.g., Koumans (1953)
noted “Nape naked before first dorsal fin”], finrays (i.e., “9?” and “12” segmented rays in second dorsal and anal fins, respectively), and teeth
rows (i.e., “Teeth in upper jaw in one row”) on
jaws. Actual meristic counts in the holotype are
given above; also, likewise the other species ofVanderhorstia, there are 3–4 rows of teeth anteriorly on each jaw in the holotype. Furthermore,according to the description and illustration madeby the previous authors, first and second spinesof first dorsal fin are prolonged in the holotype(see Fig. 4); nevertheless, actually, anterior 3
spines are greatly elongate and filamentous (third
spine longest, its length 37.0% of SL).
Arrangement of cephalic sensory-papillae
rows of the holotype (only examined specimen) of Gobius (Oxyurichthys) longimanus isquite difficult to see, and not illustrated here.
Nevertheless, the second author confirmed atleast the following features on the specimen: row

a comprising 3 papillae; row c comprises 4 papillae; anterior end of row b at mid-point between
anterior end of row a and posterior end of row c(at least on right side of head); condition of row
cp and posterior part of row b uncertain; a pair ofsensory papillae just behind chin (row f ).
In addition to these features, the other generalcharacteristics of the holotype of Gobius(Oxyurichthys) longimanus also agree well withthose of the large-scale group of Vanderhorstia.
This species, V. longimanus, is most similar tothe other deep-dwelling congener, V. puncticeps,
sharing relatively slender body (its depth 13.0–
15.1% of SL), short jaws (jaw length 12.4–12.7%
of SL), slender caudal peduncle (its depth 43.9–
47.1% of its length) and long, lanceolate caudal
fin, as well as similar meristic counts (e.g., usually VI-I, 11 dorsal-fin rays; I, 11 anal-fin rays;
19–20 pectoral-fin rays; 26–31 longitudinalscales; 8–9 transverse scales from origin of second dorsal fin downward and backward to baseof anal fin); these are also similar in their small
body size (largest known specimen, 39.9 mm
SL). Vanderhorstia longimanus is, however, readily distinguished from V. puncticeps in having:
scales on base of pectoral fin (vs. base of pectoral
fin naked in V. puncticeps); single predorsal scale(vs. none); anterior 3 spine of first dorsal fin
greatly prolonged and filamentous, extending

New Species of Vanderhorstia

posteriorly to slightly beyond base of seventhsegmented ray of second dorsal fin when adpressed (vs. spines of first dorsal fin slightly prolonged, extending posteriorly to base of first, second or third segmented ray of second dorsal fin);pale spots on head and nape more or less circular(vs. irregular shaped pale markings on cheek, operculum and nape).
Vanderhorstia puncticeps (Deng and Xiong in
Xu et al., 1980)(New Japanese name: Ho’obeni-otohime-haze)(Figs. 1D and 2B; Table 1)
Ctenogobius puncticeps Deng and Xiong in Xu et al.,
1980: 180 (original description); Cheng and Zheng,
1987: 446, 1336 (brief description in key and line
drawing).
Vanderhorstia sp. Shibukawa and Suzuki, 2004: 118 (noteon specimen; Sagami Bay, Pacific coast of Honshu,
Japan).
Material examined. BSKU 77244, 1 specimen, male,30.3 mm SL, Tosa Bay, off Kochi Prefecture of Shikoku,
Japan (33°18.22N, 133°36.08E–33°17.00N, 133°34.22
E), 120–123 m depth, 2 Feb. 2006, R/V Kotaka-maru;
BSKU 77247, 1 specimen, Tosa Bay, off Kochi Prefecture, Shikoku, Japan (33°18.42N, 133°36.28E 33°17.85
N, 133°35.82E), 120–121 m depth, 2 Aug. 2005, R/V
Kotaka-maru; NSMT-P R798, 1 specimen, male, 39.9 mm
SL, Kan’non-tsukadashi, Amadaiba, Sagami Bay, Pacificcoast of Honshu, Japan, 60 m depth, 20 July 1952.

Diagnosis. Vanderhorstia puncticeps is distinguished from the congeners in having the following combination of characters: 11–12 (usually 11) segmented rays on second dorsal fin;
26–27 (30?) longitudinal scale count; no predorsal scales; 8 transverse scales from origin of
second dorsal fin backward and downward toanal-fin base; pectoral-fin base naked; anterior 3
spines of first dorsal fin frequently elongate andfilamentous, extending posteriorly to base of
first, second or third segmented ray of second
dorsal fin; connecting membrane between innermost rays well developed; when freshly collected,
irregular shaped yellow (pale in preservative)
markings on cheek and operculum; oblique narrow yellow line from posterior end of eye to201nape; small bright yellowish red spot on center oranteroventral part of operculum; ca. 9 narrow
yellow vertical bars on body; distal tip of firstdorsal fin vivid red; narrow yellow longitudinalband on middle of anal fin.
Description. Following descriptions arechiefly based on 2 of 3 specimens examined, i.e.,
BSKU 77244 and NSMT-P R798; many characteristics of the remaining examined specimen,
BSKU 77247, are not confirmable, since thespecimen is heavily damaged. The frequency ofeach count is given in the parentheses following
relevant count. Dorsal-fin rays VI-I, 11 (2) or VI-I,
12 (1); anal-fin rays I, 11 (2); pectoral-fin rays 20(5); pelvic-fin rays I, 5 (6); segmented caudal-fin
rays 98 (2), including 77 (1) or 88 (1)branched rays; dorsal unsegmented caudal-fin
rays 7 (1) or 8 (1); ventral unsegmented caudalfin rays 7 (2); longitudinal scale count 26 (1) or
27 (3); transverse scale count from anal-fin origin
dorsoanteriorly to first dorsal-fin base 9 (2) or 10(2); transverse scale count from anal-fin origin
dorsoposteriorly to second dorsal-fin base 8 (3)or 9 (1); transverse scale count from second dorsal-fin origin ventroposteriorly to anal-fin base 8(4); predorsal scales 0 (2); circumpeduncularscales 12 (2); gill rakers 29 (1) or 39 (1);pseudobranchial filaments 8 (1) or 9 (1).
Coloration when freshly collected (based oncolor photographs of BSKU 77244, Fig. 2C).
Ground color of head grayish white tinged withpink ventrally, light orange brown or dark grayish
brown dorsally; cheek and operculum with several irregular-shaped small yellow markings, thosealong posterior part of upper jaw longest; a smallbright yellowish red spot on center of anteroventral part of operculum; iris dull yellow or bronze,
darkened dorsally; narrow oblique yellow linefrom posterior end of eye to nape; ground colorof body grayish white, slightly tinged with pink;
belly pale; about 9 narrow vertical yellow bars onside of body; a series of 4 irregularly shaped darkbrown blotches on midlateral body, the posteriormost one at middle (or slightly posterior part) of
caudal peduncle; first dorsal fin translucent,tinged with light gray, many yellow spots or yel-202

A. Iwata, K. Shibukawa and N. Ohnishi

low longitudinal line; dorsal tip of first dorsal fin
bright red; second dorsal fin translucent, tinged
with light gray (darkened dorsoposteriorly), withmany small yellow spots; anal fin white, tinged
with gray distally, with a narrow yellow longitudinal stripe at middle; caudal fin translucent,margined narrowly with yellow dorsally, with anarrow arc-shaped yellow line at anterior part, indistinct dusky blotch just behind base of the fin,several yellow elongate spots or short lines at
middle and posterior parts, and a narrow submarginal yellow band along ventral margin; dorsal
part of caudal fin narrowly margined with yellow;
middle of caudal fin with several yellow spots or
short lines; pectoral fin translucent; pelvic fin pale,with numerous minute melanophores.
Color in alcohol. Similar to color whenfreshly collected, except as follows: all yellowish,
pinkish and reddish markings on head body turnto pale; all whitish, yellowish and reddish markings of fins turn to translucent.
Distribution and habitat. The specimens
examined here were collected from off the Pacific coasts of temperate area of Japan, 60–123 m
depths. Type specimens of Ctenogobius puncticeps (Vanderhorstia puncticeps) were collected
at the depths of 76–98 m in the East China Sea,
off Wenzhou, Zheijiang Province of China.
Remarks. The examined specimens agree
well with the original description of Ctenogobius
puncticeps made by Deng and Xiong in Xu et al.
(1980). The species is clearly assigned to thelarge-scale group of Vanderhorstia by having all
diagnostic features of this group listed above.
Ctenogobius is the gobionelline genus (Pezold,
1993, 2004), known only from the Americas; it isreadily distinguished from Vanderhorstia in having, e.g., paired pore C (vs. unpaired), distinct
transverse pattern of sensory-papillae rows on
cheek (vs. longitudinal pattern), P-V 3/I II II I
0/9 (vs. 3/II II I I 0/9), 2 epurals (vs. single).
Although almost all color patterns have been
already faded, the specimen (NSMT-P R798) reported as unidentified species by Shibukawa andSuzuki (2004) is re-identified as Vanderhorstia
puncticeps in this study. The specimen, collected

Fig. 6. Dorsolateral views of heads of two
species of Vanderhorstia, showing cephalicsensory canal pores (indicated by roman uppercase letters) and trough just behind eye (indicated by arrows). A: Vanderhorstia auropunctata, KPM-NI 2791, female, 58.7 mm SL;
B: Vanderhorstia rapa, NSMT-P 73119, holotype, female, 50.3 mm SL. Photographed by
K. Shibukawa.

from Sagami Bay of Honshu, Japan, represents
the northernmost record for the species.
Vanderhorstia rapa sp. nov.
(New Japanese name: Nanohanafubuki-haze)(Figs. 1E, 2C, 4F–G and 6B; Table 1)
Vanderhorstia sp. 7: Senou et al., 2004: 367 (underwaterphotograph and short description; Kochi Prefecture,
Shikoku, Japan)
Holotype. NSMT-P 73119, 50.3 mm SL, female, off
Sankakubae, Kashiwa-jima Island, off southwest coast of
Shikoku, Japan, 37 m depth, 23 July 1997, collected by A.
Iwata.

Diagnosis. Vanderhorstia rapa is distinguished from the congeners in having the following combination of characters: 11 segmented rays
on second dorsal fin; 33–35 longitudinal scalecount; no predorsal scales; 11–12 transverse
scale count from origin of second dorsal fin

New Species of Vanderhorstia

backward and downward to anal-fin base; pectoral-fin base naked; third spine of first dorsal fin
longest, slightly elongate and filamentous; sensory-papillae row c comprising four sensory papillae; a deep trough along posterior margin of eyes
(Fig. 5B); first dorsal fin with numerous minutescattered yellow spots and no yellow submarginallines; four faint brownish (tinged with yellow)
mid-lateral blotches (rather than saddle-like bars)
on body.
Description. In the following description,
the values where we take bilaterally are separated
by a slash, the first value representing the left
count. Dorsal-fin rays VI-I, 11; anal-fin rays I,
11; pectoral-fin rays 20/21; pelvic-fin rays I, 5/I,
5; segmented caudal-fin rays 98, including 7
7 branched rays; dorsal unsegmented caudal-fin
rays 8; ventral unsegmented caudal-fin rays 8;
longitudinal scale count 33/35; transverse scalecount from anal-fin origin dorsoanteriorly to firstdorsal-fin base 16/16; transverse scale countfrom anal-fin origin dorsoposteriorly to second
dorsal-fin base 11/11; transverse scale countfrom second dorsal-fin origin ventroposteriorly toanal-fin base 12/11; predorsal scales 0; circumpeduncular scales 12; gill rakers 311/310;
pseudobranchial filaments 12/11.
Color when alive or fresh [based on color photographs of holotype, Fig. 2C, and underwaterphotographs found in, e.g., Senou et al. (2004:367)]. Ground color of head and body pale,slightly grayish dorsally; cheek, operculum, occipital region, nape and body scattered with numerous small vivid-yellow spots (smaller than ca.
one-third of pupil); 2 vertical yellow lines on
belly; suborbital area slightly darkened, with abroad yellow vertical bar; yellow line along dorsal margin of upper jaw; 4 large, faint brownish(tinged with yellow) blotches on mid-lateralbody; first blotch below first dorsal fin, second
blotch below anterior half of second dorsal fin,third blotch on anteriormost part of caudal peduncle, and fourth and posteriormost blotch at
caudal-fin base; 4 indistinct, shallow saddles onbody; first saddle around posterior half of firstdorsal-fin base, second saddle at middle of sec-203

ond dorsal-fin base, third saddle just behind posterior end of second dorsal fin, and fourth andposteriormost saddle just above lateral blotch at
caudal-fin base; first dorsal fin translucent andslightly whitish, with ca. 8–9 irregular longitudinal rows of small yellow spots; second dorsal fintranslucent and slightly whitish, with 3 longitudinal rows of small yellow spots (spots on distal
row larger than those on proximal two rows);
anal fin slightly whitish, a little tinged with blue
distally; soft anal-fin rays tinged with yellow;
caudal fin translucent and slightly whitish, withmany elongate yellow spots on middle and upper
part; rays of ventral half of caudal fin tinged withyellow; fin membranes of lower part of caudal fin
a little tinged with blue; pectoral fin translucent;pelvic fin pale white.
Color in alcohol. Similar to live coloration,
except as follows: ground color of head and body
pale yellow or pale brown, becoming darkened
dorsally; all yellow spots on head, body and finspale (these spots on body quite indistinct, exclusive of those on dusky areas); bluish areas of finsdusky; iris entirely blackish.
Distribution and habitat. Vanderhorstia
rapa is described based on a single specimen
from Kashiwa-jima Island, off southwest coast of
Shikoku, Japan. It is found in the coarse sandy
bottom around rubble area (25–50 m depths) in
Kochi Prefecture of Shikoku, and symbioticallyassociates with the alpheid shrimps (Senou et al.,2004). A specimen (Fig. 4H), photographed at
the depth of 39 m in Izu-oshima Island of Izu Islands, is provisionally identified as V. rapa, judging from its coloration of head, body and fins,and squamation.
Etymology. The new species is named Vanderhorstia rapa (Latin rapum, meaning “turnip”
or “rape”), comparing its minute golden yellow
spots on head, body and fins to the rape blossoms.
Remarks. Vanderhorstia rapa quite resembles V. auropunctata in general physiognomy, andwe have once considered that the former might
be possibly the intra-specific color variant of thelatter. However, subsequent detail examination

204

A. Iwata, K. Shibukawa and N. Ohnishi

based on the actual specimens reveals that these
2 are distinguished in size of jaws (jaw length
50.8% of head length in V. rapa vs. 43.9–44.1%
in V. auropunctata), shape of trough along posterior margin of eyes (distinct and deep in V. rapa
vs. indistinct and shallow in V. auropunctata),
elongation pattern of spines of first dorsal fin(third spine longest in V. rapa vs. fourth spine
subequal or longer than third spine in V. auropunctata), and number of sensory papillae on row
c (4 in V. rapa vs. 5 in V. auropunctata), as well
as discrepancies in coloration of fins and body.
Because all examined specimens of these species
are females, it appears to suggest that these differences are not linked to gender.
Acknowledgments
We express our sincere thanks to the following
persons and institutions for specimen loans
and/or registrations, assistance during the visit totheir institutions, help in field collection, andproviding various information: H. Endo and E.
Katayama (BSKU); W. Hiramatsu (Ehime Prefecture, Japan); H. Senou (KPM); K. Matsuura
and G. Shinohara (NSMT); T. Hirata (Kochi Prefecture, Japan); R. Vonk (ZMA). T. Hirata and Y.
Miyamoto (Kanagawa Prefecture, Japan) kindly
provided the underwater photographs. This study
was partly supported by the Japan Ministry of
Education, Science, Sports and Culture Grant-inAid for Scientific Research on Priority Areas(#319), Project “Symbiotic Biosphere: An Ecological Interaction Network Promoting the Coexistence of Many Species.”
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Manuscript received 15 March 2006; revised 28 December 2006; accepted 10 January 2007.
Associate editor: K. Matsuura.